Khurana, Subhash Chander (1981) Growth, development and...

286
Khurana, Subhash Chander (1981) Growth, development and yield of potatoes. PhD thesis, University of Nottingham. Access from the University of Nottingham repository: http://eprints.nottingham.ac.uk/13729/1/256347.pdf Copyright and reuse: The Nottingham ePrints service makes this work by researchers of the University of Nottingham available open access under the following conditions. This article is made available under the University of Nottingham End User licence and may be reused according to the conditions of the licence. For more details see: http://eprints.nottingham.ac.uk/end_user_agreement.pdf For more information, please contact [email protected]

Transcript of Khurana, Subhash Chander (1981) Growth, development and...

Page 1: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

Khurana, Subhash Chander (1981) Growth, development and yield of potatoes. PhD thesis, University of Nottingham.

Access from the University of Nottingham repository: http://eprints.nottingham.ac.uk/13729/1/256347.pdf

Copyright and reuse:

The Nottingham ePrints service makes this work by researchers of the University of Nottingham available open access under the following conditions.

This article is made available under the University of Nottingham End User licence and may be reused according to the conditions of the licence. For more details see: http://eprints.nottingham.ac.uk/end_user_agreement.pdf

For more information, please contact [email protected]

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GHOid'l'H 2 DEVi~I..DPHENrr fJJD YIELD

OF POTATOZS

by

SlJPTj".C::;.l CF!"Dnn TTIIUnA····1A p S (P P U TUDllIA··IA)_Jil'._"J_ .[1-\1') l,jl\ 1\.. .Li. ~~ , .L.J. C. ...,•. \.•• ,.LJ il 1~

F.Sc. (Il.A.U.,EISSAP).

Thesis submitted to the

Un i ver-si ty of Nottingham for the degree of

Doctor of Philosophy

Dep)rtmeLt of Agriculture and HorticultureUn i versi ty of No t t i ngham

School of AGricultureSutton Bonington

LoughboroughLeics.

IJ ovember, 1981

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BEST COpy

AVAILABLETEXT IN ORIGINAL ISCLOSE TO THE EDGE OFTHE PAGE

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BEST COpy..

, .AVAILABLE

, Var'iable print quality

,

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TO MY PARENTS

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ACKNOWLEDGEMENTS

I am greatly indebted to Professor J. D. Ivins and

Dr. J. S. McLaren for their interest, enthusiastic guidance

and encouragement throughout the present work.

I would also like to thank Mr. J. Craigen and other

members of the Biometry unit, Library staff, Mr. C. Roberts,

all technical staff of the department of Agriculture and

Horticulture, especiallY Mr. D. Hodson and Mrs B. Hull and all

others who helped in the present work.

Finally I wish to express my thanks to the Association

of Commonwealth Universities for financial assistance and

Haryana Agricultural University for study leave.

S.C.K.

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CO::T;';r··TS

1 IJ:TRODuc'rIO;:

P~'be

1

2.1 3

2.1.1 Effects of pho tope r i od2.1.2 Effects of temper" ture2.1.:~) Effec LE;of liC!lt

345

2.2 om l~CTIVL3 7

2.3 8

2.3.1 Exneriment GR1 8

2.3.1.1 Prenaro.tion of pots and p.lan t i ng2.3.1.2 'I'r e a trne n t s2.3.1.3 Exoe r imen tu l desiGn 'lEd pr-ac t i ca'l

detn.ils2.3.1.1+ Gr-ow th c:lr..Alysis

2.3.2 Ex'.)erimer.t GR2

2.3.2.1 Pr-epar at i.on of pots and p lan ting2.3.2.2 Treatments2.3.2.3 Experimental design and pr-ac t.Lco.l

details2.3.2.4 Gr owth ilu,dysi.s

2.3.3 Experiment GH1

2.3.3.1 Pre ~nra t i on of :)01;S .md p.lant ing2.3.3.2 '1'1'e ,_l tme n ts2.3.3.3 Experimental design and pr-ac t i.cal,

details2.3.3.1+ Gr-owth .::U1 ::.J.l ys is

2.4 m~SUL'rs

B8

99

10

1011

1111

12

1212

1213

15

2.4.1 Exoe r iment Gl{1 15

2.1+.1.12.4.1.22.1+.1.32 .li. 1 .1+2.4.1.5

StolonStem

1515151617

Le;_,fTuber':::'otal dry mat tel' .ic cumul a t ion

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19

2.1+.2.1 Stolon2.1~.c:.2 Stem2.L:-.2.3Le:::2.h.2.1+ Tuber?l:-.2.5 ToLil rlr,Y r:_d:tol' -:ccu::mlJ1_jo,

2.4·.3.12.1+.3.22.4.3.32. i+.3. 1+2.1~.3.5

DISCUSSIOJ;

1920202122

23StOJ_OllStem1,0 '.fTuber'I'o ta.I dry Height

232323242426

3 CHEHICAL co::r~ROL OF GHO'.vTH

3.3

3.4

3.5

osr [_;CTIV1~~S

3133

j:AT~mIALS Mm HETlIODS 34

3.3.23.3.3

3.ll-.13.4.23.1+.33.ll-.43.4.5DISCUSSIo;~

Expe r i.men t.a.l design and pr ac t i cu Lde t.ri LsGrowth :m·J.]_ysisFin_,_l harvesting

34363639

Stolon and stem growthLeaf grow thLigh t .in 'te r-ce p t i.onTube r c;rowth ,.And development'I'o t.rl. dry mc.t te r accurnul.a t i.on

394041424344

4.1

lj.2

4.1.1il-.1.24.1.3

Phys.i o.l og i c al, i,\ge1:1 t.e r=-o r-opp i.ngLicht Ln te r-cept ion

4949525355

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/. "Zt • ./

Exner imer. t }'1 5'1

~·.j.1.1 Exr.er i.mentzL desiGn ,~'ld r)r;).ctic "1

de t ..:i.~~sIf.3.1.2L~.3. 1 .34 •.5.1.44.3.1.5

4.3.24.3.2.1

1+.3.2.2~.3.2.34.3.2.[1-4.3.2.5

4.3.3.2It.3.3.31+.3.3.4

Lt. 3. 1+. 24.3.4.3Lt.:5.t~.4

/+.3.5.24.3.5.3,

4.3.6.211-.3.6.3

G:{'O\'Jth ;:u.-~r.J_ysisSpr-cu t ::;rov!thduriES stort\geIle.rsur-emen t of soil ':i',tercon te : L

5,'758595960

,C;xperiment F2 60Expe r irnen tal design Qndpr:_;_cticalde taiLaGrowth ana,lysisSprout growth during storageMeasurement of soil water cortentFinal harvesting

606162626263

Experimental design and or-ac t i c.d.de t.ri LsG r 0\" th.m :.llysisSprout ~rowth duriGG storageFi:1n.l hur vestir:l~

Experimental design ~nd practic.Qde tai LsGr-ow th clTlalysisSprout growth during storuGeF'inul hurves t iug

65666666

Experiment F5 67

Exper-Lme n t.al. design and pr ac t i cc.IdebilsGrowth :..m:uysisS)rottt growth during stori..lge

676868

Experiment F6 68Experimental design and oracticalde t.ri LsGr-owth a,n:llysisSJrout growth during storage

686969

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I ,-'+._;

1+.1.;..2.1.1+. Lf-. 2. 2.Ir.lr.2.3

4.1.1.2-"4·.4.2.5-1+. Lt. ?(;

Lt. !+. 3.1i!.Lf-.3.2It. Lt. 3. 3

Lr. 4. 3. Lf11.i.;.3.5LI.L!.3.6

4.4.6.1

L~.i+.~.1!1.L1-.7.2

'1.h.? 34. L~.'? .1+

1+.4. (4

DISCUSSIO~

If-.5.1L1-.5.2£+.5.31+.5.l1

:=xnerirnent }'1

E:;lerG81~ce :.c:d stem nurce rSt02.0L grov!th nr:d de ve Loprne rtGr ow th ,·'.nd de ve Lopme nt of s tem.I'dl erfIJigh t b ter-ce ot ionTuber grO'.·:th and development'I'ot aI dry r;,,:;tter .iccumul.c t ion

Experiments: F2; F3; F4':~mercellce .uid stem numberStolon growth u~d developmentGr-ow th .u.d development of stemTid LeifLight interceptionTuber gr0\1th).lld development'I'o t al. dry mu t te r .iccumul.ct i on

Conroe ti t i on be twe e r; two compone n tsw i th in plot

Crop ev~por~tion

Light interception und pot~to growth

Leaf ~rea index ~nd ohotosyntheticallv.c t i vo r,.ldLltion (PAIn interception ~Jry matter oroduction

Ex.ie r i.rne n t FS

Emergence uid stem numberGrovlth '.Dj de vel.opmen t of stemi..i.:d ledTuber growth and development'_L'otD dry m:,tter .ic cumu.Lat ion

3~rout gro~th during storageEr:lerc;enceTuber Cro\Jth .u.d de veLopmerrtGe~erul crop gro~th

'70

'70

81

8132

83868689909094

959898

102

103

106

108

108109

111

111

112113114

114

118

118120122127

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5

6 13/

'? AP?Ei:DICEL3

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GEC.-JrE I Al·:D

AJSTFACr

,~x0er:~rrel~ ts v.ore CO:lC;UC Led L. i~ro"J~;l r oor.s LO otudj t~e oh o to-

ne r i odi c r-e scouse o~' Fer. ::~. Lei Crown. 11, f ietd exoe r ime nt s the effects

of s)routi,[_; l.ech r: -ue s :..ul the ')o.sslbi.ity o f nii x i rg of d i f'f'e r-e . L

'Jbysiologj,c,d,l:y,,-?;ed tubers dcdl use O[.';.)l: t gr-ow th reGu11 tor I -)0333

»ies:« studied. [))rout JrO'llth of the tubers used for :)l:,ntillg 'Jos studied

dur i:1C s tor i;e. ~"ight .in Lor-c e o t Lon ~ LLc ohot ocyn the t i.cul.t.y act i ve

i t :.':cl'e:!sed ttcJ.oc l t i.or. of ,"i.:3simiL; te a to tte tubers, wh i.ch resulted

Sprout ;:;ro·,.-lLh \\",S l ine ar ly r-e , .t ed to the .i n i t ieL t ube r we i gh t and

ri <'l degrees . 4°~ r 1.co ve C' rom o o i-m.mcy St()ri'~ tubers in the cold

Le fore nl·J. ti ~>'; increlse:i the w:ri),ui nur.oo r ,:11 thus the pr opor t ior; of

lYlin s tems i· t h e f i.eLd , Iirne to reach 50,.'; en.e r-ge nce \'Id,S not reduced

"'1 ( t '+10" , .. , • I' "t')'~o.d tubers stored .', ;J _ l, un CLL .t ell;_,' oe ore [l.L,(Jl lng delayed

tuber ini t irt i o.. out or.ce the tuber 11:d oe e n id, tiiAted the n the effect

growth or developme~t ~.s ~o more ~ffected by the physiological age

of the seed tuber. YieLls from m.ix i.ug different phye i.o.Lcg i co.Ll.y aged

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tubers were rot different from those expected.

1'1;e DL.jor f.!ctordfecting the gro ..!th of t.ie cr-op 1I"S the 'd.,ter

su~')ly. ':_'heo'Jer~:;_l "hotosynthetic conve r-a ior: e f'f'ec iency of the

( d . , t v T -1 ',"" '.t- I' ) I.~,g . r,Y \'Jelgn!"lt.. ':'.~, l:J cer ce o teo i'i .ie 3. '+23 ar: 1,)(,:') in ,ose nce

c~nopy

of

,.LIlY app.rr-e r.t \·.':.tel' stress whi I.e il 1(),/S; .i t ,LS cnl.y 2.L;9c due to w.ite r

stress.

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1

Crop C:,LOOY is b,sic·.l·:~Y " cor.ve rte r of soLar ener-gy to dry matter.

Po t.z to crop ~ro':Jth h.rs oee r; considered or..Ly in terms of led ar-e.; index

(LAI), 1e"f,ro' durtiol~:.ud :leLlSSir.lj~.~tioL r-ite , Cr-op gr-owth r.a te

for var ious .Jgricuj t.urc.I crops h.rs been f'ourd to be pos i t iveLy corre-

lilted to the liGht .inte r-cep t.ed by the c:;r:o:)/ (Biscoe .ind GCJ..Llclgher,

19'1'7; ·,'.liEi3ms et al., 1965). Published r-e nor ts on 1igh t iL terce ptionc

in _l)ot'ltoesare f'ew (Scott and i;iilco}{son, 1978).~

There 'iPge~r to be i.l. considerable genetic difference in the photo-

periodic dependence of po ta toe s (Hur t i et :.1., 1975; 1"lendo<:;aand Hayne s ,

19'(6). Potutoe a wre o.l nnted .i t different times in the field due to

various r-exsons such :{S wea the r, It is accented 'tha t there is .;l. oal ance

be tween growth of tubers and rest of the plant, .myth i ng whi.ch favours

the growth of one w iLl retard the gr owth of others (Eoorliy. 1978;

Varieties differ in their rutes of sprout growth at Lt given temp-

erature (Headf'o rd , 1962; Short :md Shotton, 1970). Physiologically old

seed h.xs beer: found to Lncr-e aae tuber yield ear Ly in the season but the

effect change d :J.G the hrr ves t i r.g Wi.'S delayed and in some cuse a it became

negative (0'3rien ~Ld Allen, 1978; Allen et al., 1979) which was assoc-

ia. ted w i th Lowe r L1\1 Ln the caae of old seed, whi ch may be overcome by

m·iEipuL.i.ting ap.ic ing , Higher tuber yields have been reported by mixing

eo.rly"u'1.d Late crop var ieties (Schepers .md Sibma" 1976), but the mixed

product m~y be o~ly useful for st3rch industry or other uses for which

~ mixed product is ~cceptable.

In this the s i s r esponse of Pen t l and Crown to photoperiod and irr-

:..tdLJ:lce is studied it: Growth rooms. Effec ts ofnhysiologic"ll age,

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2

t.ube r-scu.d 1}r:J333 (:):;_ ..uit f,rO'., t:l reguJ."tor, rci: :1,3 be e r, s tud.i e d in the

field by aocuen t i c.I har-ves ts for cr-o» gro't,th ;.Iul,}'sisllollg w i th regular

1 igh L measur eme II ts.

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2. PHOTOPERIOD AND LIGHT

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2.1 LI'l':~Ef1TUIE l';_~VIc.::,l

l~ffects of envi r-onrner: t h ..ive lone; oee.. reeosnised 'JS 'Ur:ODe; the most

i:;:~ort:(nt f:~cton' Lnf'Lue nc ing tuber f'orrn.i t i.o.: by the~lot"to 81il.nt

(G~ir:'Jer:1l1d AI1(rd, 1923; '3u<.;!l:1811, 1925; rcClel1..,lfd, 192E; Dor-oshe nko

et 'll., 1930; /,rthur et,1.., 1930; "ier:wr, 1940; Driver .uid Huwke s ,

19L1-3). 'I'he re is~c~:no.st:;eu::r'{l .'\_;reei:lent tl.o t env i r-onrne n ta l factors

wni ch s t irnulate hau i.ri ~ro1:lthJ-=-so deLay or i,:hibi t tube r i z.c t ion ,

2.1.1 Effects of nhoto~eriod

'l'here'C~~)e,r to be cons i de r-abt e genetic differences .ir: the photo-

period de oer.dence of l)obtoes. Schick (1931) observed that four German

vur i.e t i es showed 1i t tle r-eaponae to a reduction of pho tope r iods to 12

hours, wh~le three South Americ~l varieties showed a very striking

response. Different resoonse to photoperiod, for the varieties Alpha

.nd Eer-steL'ing h:)5 been reported by Bodl aende r and Earinus (1969), arid

l<urti et 3,1., (1975) r-eoor ted for the varieties Kufri Lauvl ar , Kufri

Jyoti, Kufri Jiev~'-r:.,nd SLB/~,~405 a. Mendoza ar.d Huyne s (1976) re-

~orted differerces i!1 critical day length ~mong potuto clones of three

Llxonomic gr oups r and i.ge r.a ;phure jc;; tuberosum.

The vur i e ty Kuf'r i, Sindhuri, previously grown e,t oon t inuouaT igh t

for 47 d~ys did not form tubers for 60 d~ys ~t 14 hours day length while

cn.l y 11 short d~Jys (g hours) were required for tubers to initiate (Hurti

and Bane r jee , 1976:J.).

l<urti et.\l., (1975) reported Lncr e.iso in tuber yields in var i e t ies

Kufri Lauvk ar , Kufri Jyoti, Kufri .Jee vun arid SLB/Z 405,-, under short days

cornpar-ed to nu tur-u l dDY length in summer (temperate c.li.mate },

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IIigher tuber y i eLde ur.der short diys (e,-1 () hours) compar-ed to long

d ays (18 hours) lnv8l1so beer: r-e norte d iJ,Y some other wo.rke r-s , GreGory

::'nd Chapm..n (1962) fa!' ned ;';cClure. Lr.cont r-rst to tl.e se , aome wor-ker-a

h uve r-epor te d highe r yields unde r 1011[,; d'q.s (18 hours) compar-ed to

short days (12 hours) t Bo.r.m (1 ~)59) for the v-u-Le ty Arr:d: Pilotmd

:;:.odJ.u,ender .md Ii;urinus~?o~ ~erstelir:g and 1\.1;;1h:lo

Iricr-ease in tuber .induc bon due to pho tope r-Lodi sm h.is been r-eLated

to Lr.cre ase in :1eve Is of c,'!tokio I r;s (L'ngilLe ;,nd Forsline, 1974 j Forsline

and Langille, 1975), and de cr euae in levels of gibberel1ins (Okazawa,

1960 j Pail ton v.nd ',Jale ing, 1973), w i th increase in short day cycles.

2.1.2 Effects of temoer~ture

The effect of tempe r-a tur-e is s imiLar to th.rt of photoperiod, higher

temperature promoting vegetative growth wld ~ower temperature stimulat-

ing tube r-Lza t ion , P'lan t s of the variety Sebago previously grown under

non .induc iug coudi t ione g:we higher tuber yields ':At 22/18 compared

to 32/1SoC (r'iellzel, 1930). Bor-ah and I'Hlthor~e (1962) reported decrease

in the oercentQge of assimilates diverted to the tubers with increase

in terr:per.:J,ture:.;,nd Gregory (1954) reported .et decrease in tuber to top

weight ratio with increase in temper~ture. Tuber initiation was in-duced by Lowe r i rig the t.emoer a tur-e to '/°C for about a week , in plants

growing at 20/15 or 25/15°C (16 hours day length) by Burt (1964).

Sah a et al. ,(197ll') r-enor ted an incre.').se in tuber yield \,..ith de cr ease

in only night temperature. Like photoperiod there is considerable

var i ance among pot.i to varieties .in t.eraper-ctur-e dependence. i:Jith in-

cre'.lse in night temper:::.ture from 20 to 30oC, reduction in tuber yield

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was 80 and '75;0 for the v.ir i e ties Kuf r i Jyoti and Kuf r ; Chandr-amukh i

respecti vely, while Kuf'ri Sindhuri Lliled to form tubers at 300C (Saba

et i{l., 1974).

Optimum t.emper-ut.ur-e requirement for cell;)' var i.e ty is influenced by

other envi r-onmea ta.l L~ctorG such as ,1hotoperiod and irr:)di::.Ylce.

vr,s 8 hours .uid ~:t 17/1UoC where day length w:~s 16 hours.

Effects of light

The influence of light on growth and yield of potatoes is deter-

mined by the irn.dicuJce level, its duration and qua.l i ty of the ligh t ,

7'hotosYllthesis of :J si:l;:;le Le uf :is wel.L :18 of the CLUJOpyQS ;:, whole

Lncr-e ase s Viith .inc rease in Lr-r ad iance until an optimum is reached

(Ku et al., 19?7j Sale, 1974). Pohjuhk.a.lLi o (1951) reported change in

top to tuber r-a t i o HI f'uvour of top with decrease in Lr r-adi ance , Borah

(1959), r-epor ted a 68 to '7?;<: decrease in tuber dry wei gh t with decrease

in irradiance by 45;',; in the var i.e ty Ar-r-an Pilot, in both the green

house and field. Differences were higher .ir: growth rooms, tuber weight

" -2-1per pl an t ut 64 .md 120 ca l. cm d Wb.S 1 and 14g respectively.

SimiL~r resul t s were r e oor t.e d by BodLaonda r (1963), in this experiment

tuber ini t i a tion "Ivas deLaye d by 27 and 54 days vii th de cr ease in irradi-

Cince from 16000 to 8000 'md 3000 lux respectively. Even in a region of

high so lar input (d;-,ily soLtr input .iver age s 25-30 ~W:ln-2), Sale (19'73)

r'epor ted de cr ease in tuber number- :u;d increase in time between tuber

ini t ia tion arid rnax i.mumbul.k i.ng rette, w i th increase in shading.

Exper Lments unde r con trolled condi t i oris provide very useful

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6

Lnf'or mat i.on , but s oe c t r a.l photon distribution (SPD) tuy af'f'e c t several

as oe c t s of »Lan t gr owth and de veLoomont (hcL~1.reIl .ind Smith, 1978) and- -

SPD is different for di I'f'e r-en t light sources (!.icCree, 19'7c~).

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2.2

There·u·e cona Lder-able differences in response of various po t« to

varieties to the euv i r-onmer.t (Ch.ip te r 2.1 ).Dot·:.toes :{re »Lan ted at

different times .').nd so some ::)hoto_oeriociicnd lizht t r-ea trnents wer-e

given to the vur i e ty Pen tLand Cr-own La unde r-s t.rnd its response to the

env i r-onmen t .JS this v.ir i e ty \~:l.S to be used .in 'I number of experiments

in the field.

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"u

2.3 i'lA'rERIALS Mm NE'l'HODS

Out of three, two (Expe r irnenta : GR1 ar.d GR2) were carried out in

growth rooms and third one in g.lasehouse (;::xperiment GB1). Potu to

var i.e ty Pen tLand Crown "/iAS used for '-1.11the experiments.

2.3.1 Experiment GE1

Preparation of oats and p1~nting

Pots of 25cm diameter wer-e washed thoroughly in hot water and

filled with John Innes potting compost No.1 (de tu iLs given in Appendix A).

One mmnyl.on rnesh was kept over the compost and apically sprouted

tubers (details for seed source given in Appendix B) of uniform size

were p13nted on top of the mesh to study tuber initiation and stolon

growth per i.odi cul Ly , Pots (indi vi duul.Ly ) were. covered with double

polythene shee t whi te inside arid black outside, wi th a hole for sprout

to come a\Jt. Pots were kept in gr owth rooms at cons t an t temperature

of 15°C and 16hrs photoperiod until emergence.

Tre Cl tmen ts

After em~rgence on 12 Feb. 1979 pots were given photoperiodic

treatment according to 'I'ubl e 2.3.1.1. In growth rooms (Saxhill),

warm whi te fluorescent tubes were used as the source of light.

Irr:.J.di::wce in pho tosyn the t i cel Ly ac t i ve r:lJ1ge (400 - 700nm) was main-

t.J.ined:"s 450 UE~-2S-1 (Photon fluence r:.:tte) on pot level (measured

Using qUClntummeters with cosine corrected sensor heddsj lambda

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)

instrument). Photoperiod: 16 hours day in case of long day (LD) and

10 hours day in case of short dUJ (SD) i:<ndday-night temperatures of

200/15°C was maintained. The temperature in the growth rooms were

continuously recorded on thermographs, fluctuations being of the order

of i10C.

Ex-oerimental design and uractical details

Due to limitation in the number of growth rooms available, plants

were treated as replicates and data vias analysed as completely random-

ized design. Number of replicates used at different time are given

in Table 2.3.1.2. Although there were 1-3 SEDs, depending upon the

number of replications for different treatments (Table 2.3.1.2), but

for aimp.li.f icat ion only one SED is shown which is for comparing the

treatments with different number of replications.

After roots had grown to the bottom of the pot j 150ml of supplem-

entary solution containing: 50g (NHLj-)2S04,9.2g KfW3 and 7.5g NH4H2P04

per litre was given every week (Burt, 1964). Plants were irrigated

w i th ordinary t3.Pwater as aridwhen required. Datilon length of

stolons, le:J.vesof axillary branches and number of: stolons, leaves,

axillary branches and tubers were recorded without harvesting. Tuber

was judged when tip was swollen, twice the diameter of the actual

.stolon.

Grm.,.thAnillysis

Due to fewer number of p.lant.s ava i.Lab.l e cn.Ly2 growth analyseswere carried out: h9:lnd 66 doss after emergence.

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10

Plants were stored in a cold room (4°C) until they could be

dissected. 'I'he number of Le ave s per _?lcmt on main stem and axillary

branches were counted seDarately, at the same time, the leaflets were

removed from the pe t ioLe, Length of the rnaan stem and the'lXillary

branche s meaeur-ed arid petiole and stems were combined for stem's dry

weight. 'I'he leaf urea was determined using the DUDChmethod,

("\'i::J. tsar. and 'datson, 1953) as modified by Radley (1963), which

invol ves tak ing discs of known diameter and dr-y weight ',I[dSused

jnstead of fresh.

Fumber of maL1:lnd br-unched (defined in Apne£x E) stolons, were

removed, counted and their tot:Jl length measured. Roots were difficult

to separate from the peat and were not collected .::.tedroots which were

uttached to the stem and stolons were removed emd discarded. Tubers

were removed, riddled, counted and weighed before being sliced finely

wi th a kni f'e and dried. Dry weights wer'e obt ai.ned by drying different

parts se oar-a te Ly in the oven at 85°C to cons tan t weight.

Experiment GR2

2.3.2.1 Prepo.rution of DotS und planting

Same as described e~rlier, 2.3.1.1. except that tuber pijces

.weighing 7g (instea.d of whole tubers) w i th single eye were taken

using et cork borer (same dinme te r in .dL the c.::;.ses) and :planted in

moist aand 2cm deep at 15°C (uet:1.ils for source of seed given in,

Appendix C). After one wee.: tuber pieces were selected for uniform-

ity of spr ou t and planted on to'!') of the mesh and covered with verm-

iculite before covering with double rolythe~e sheet.

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2.3.2.2 'rre :.'tmen ts

This experiment W:lS conduc te d dur ing 1980 .uid there «ere 4 tre: ..t-

ments: oomoi ra t ions of 2 »ho tc per i ode i ,«, 16 hours (LD) .nd 10 hours

( D) 2' " 1 1 ( t - J)." Ir\~ U"'J-2S-1 (HI)S and l.rr,lQli'.Ece Leveis on po .ieve r 1.e. -oo _',j, and

290 UEl~-2S-1 (LI) (400 - ,?OOnm). Cous t.urt temoer-at ur-e of 15 ~ 1°C was

mai n t afr.e d , Res t s ame ,tS de.scri bed e:J.rlier, 2.3.1.2.

Ex~eriment~l desiGn :Ind Dr~cticul det~ils

S::uneis described e ur-Lie r , 2.3.1.3 but no d,t:J. I-U.tS recorded with-

out growth ."',n:,::'ysis exce o t tuber .ini tittiol~Uld he i gh t of the mai n stem.

Five :pJ.':"l.ntsper tr-eu trnen t wer-e used for meusur ing height of the mai,n

stem. 5, '1,3,4, .rid 3 plan t s wer-e har-ve s t ed for gro\rlth Lcllalyses carried

out c..fter 29, !~1, 51, 66 and 81 di,Ws of emergence r-es oec t i veLy ,

2.3.2.4

Same ;J.,S described ear-Li.er, 2.3.1.4, but growth :.;Jl:.J.lyses were carried

out at frequent intervals.

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12

Experiment GE1

Preparation of pots ~1d pLantl~S

Same as described earlier, 2.3.2.1, except th~t seed pieces

(2.3.2.1) were pLillted i:; 17.5cm po ts , 2cm deep if. J.I. po t t ing

comoost number 1. Fats ':fere kept i:' :;reclJ house wher e temper a tur-e

var ied from 100C min. to zo?c mu. Pots ',Jere not covered w i th any

poLy the ne sheet. Deta i.L for source of seed is given in Appendix C.

Tre;:;_tments

10 days after emergence (on, 11.4.1980), pl(mts were selected

for uniformi ty and wer-e ei ther left under na tur al, gl asehouse light

or under, muslin, red or blue shude s (Cinemoid filters vere used for

red and blue shades) in .m unhe.i te d gL,sshouse. Air maxi.mumand

minimum temper-atur-e cbtai.ned from nearby meteorological station is

gi ven in Figure 3.3. 2b. 'l'r-anam.i t tance ('+00 - '700nm) measured (2.3.1.2.)

through the sh.ides was 21.97, 33.10 and 32.67, percent for red, blue

and muslin shades respectively. Spectr;ll photon distribution measur-

ed using scanning spectroradiometer is given in Fig. 2.3.3.1.

Phytochrome state was 0.56, 0.49 and O.ltlt for muslin, red and blue

.shades r-espe c t i ve2.y and 0.54 for na tur-a l [;l:l.sshouse.

Experimenbl design il.'1dpr:Actic:,l debils

Due to number of Li.mi tc t ions of shude s , :.\vdil·.~ble, pLmts were

used as replico.tes and duta Was analysed as completely randomized

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design and there were three r-ep.l i cut es , Only mai,n stem length was

measured w i thout harve s t ing, Rest same aa described ear-Li.er , 2.3.1.3.

Grm/th :malvsis

Same :AS described e.rr-Li e r , 2.3.1.4" but only one growth analysis

\'/as curried out.

Table 2.3.1.1 ..

Det~ils of treatments

'I'r-e a trner. tname Detail

LD Long d.xys f'r-orn emergence onward.

LS 17 long ddys from emergence onward,followed by short days.

LSL 1? long days from emergence onward,f'oLl.owed by 20 short days dnd thenlong days only.

SD Short duys from emergence onward.

SL1 17 short days from emergence onward,followed by Long days.

SL2 37 short days from emergence onwar-d,f'ol l ov....ed by long days.

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rrilble 2.3.1.2. Numbe r of r-ep.l.i.c at i ons for d i f'f'e r-en t treatments used

for anal.ys icg ddt', ,it di f'f'ere r.t time.

Days afteremerc;ence

10,13and 18

23, 28,31 and 35 45 66

Treatment

LD 17 5 5 2 2

LS 12 Q 3 5u

LSL It 2 2

3D 18 12 6 2 3

SJ...1 6 6 3 3

8L2 6 4 2

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.20 .20T-

I

~ .00 .003Bo 520 660 Boo 3Bo 520 520 660 Boo

T-

I

III

C\JI

S 1.20 (c) 1.20 (d)~..~Cl) 1. 00 1.00

.Bo .Bo

1.20

1.00

.so

1.20

1.00

.60

.40

.6.0

..00

.40

.20

.003Bo 660 Boo520

Wavelength, nm

.Bo

(b)

.60

.40

.60

.40

.20

3Bo 660 Boo520

Figure 2.3.3.1 Spectral scan (400-BOOnm): (a), glasshouse (12=00);(b), red shade (12=40); (c), blue shade (12=30); (d), muslin shade(12=42), taken on 4th March 19BO, with a scanning spectroradiometer.

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2.4 IESULTS

2.4.1 Ex-oeriment GR 1

2.4.1.1 Stolol1

Length of the m.ri n and branch stolons (Append ix E) and thei.rnumber

is given in Figures 2.4.1.1. and 2.4.1.2. Stolo!:!numberet!1d their length

was not affected by the treatments until tuber initiation after which

LD continued to produce more; main u.swell 3S branch stolons and their

length was also more. Plants whi ch hud 17 short days in the beginn-

ing and then moved to LD (trei.!tment,SL1) behaved like treatment LD

and tree<tments LS and LSL l.ike SD. Data for stolon weight is given in

Table 2.4.1.2 and was higher in treatments LD and SL1.

2.4.1.2 Stem

Length of the main stem (data not presented) was not affected

until few days :lfter tuber initiation, after which pl~nts in SD started

diverting most of their assimilates to the developing tubers while in

LD stem length was still increasing. Ax iL'lar-ybranches, which started

developing little before tuber initiation (Fig. 2.4.1.3) continued to

develop and gr-ow at f aater rate in tr-eatmen ta LD and SL1, while in

others rate was very 10'01. Data for total stem weight is given in

1'able 2.4.1.1 which w';tshigher in t.r-eatmen ts LD and SL1.

2.4.1.3 Leaf

Due to shortage of number of p.lants for growth ana.lysi s it was

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A'IU

decided to count number of Leave s and measure their length without

harvesting. Internode length was not affected by any of the treat-

ments thus number of leaves and their total length did not vary until

tuber initiation afte r which LD ol an ts had more Le.ive s and their total

length was also more. Number of leaves on uxi I l ar-y br anchea were more

in treatments LD and SL1 and their to t al, length was aLso more (Fig.2.4.1.4).

Lower leaves from trea trnents: LD and 5L1 started coming off after about

40 days of emergence, but they were still growing, while in others it

started slightly later. Total leaf 'J.rea after 49 and 66 days of emerg-

ence is given in Table 2.4.1.1. Specific leaf area and ratio of leaves

to stem is also given in Table 2.4.1.1, and was lower in treatments LD

and SL1. Thus leaves in theBe treatments were either thick or stored

carbohydrates which were beillg diverted to tubers after tuber initiat-

ion in other treatments.

2.4.1.4 Tuber

_Percentage of plants forming tubers is given in Figure 2.4.1.5.

p'Lmts which were only under LD did not initiate tubers until 66 days

after emergence when the experiment ;-JaS terminated, while all pl.DB ts in

case where they had only SD or given SD after 1'.1days of emergence

formed tubers by the end of experimer: t, suggesting th.rt tuber stimulat-

ing conditions have more effect if given few days after emergence,

ro.ther than straight from emergence onwur-d, Data on number of tubers

per p'lan t is presented in Figure 2.4.1.5 and weight of tubers in Table

2.4.1.2. Although plants which were shifted to SD after 1'7 days of LD,

initiuted tuber but stimulus in the begining wus not enough, as they

had lower numbe r of tubers, but by the end of experiment (66 days after

emergence) they did not vary from the plants which were only under BD.

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1/

2.4.1.5 1'ot:J.ldry matter accumulatior:

It was observed t.h.i t roots »iere more in tre.i trne nts 1D and S11,

but could not be se o.sr-a ted, thus to txl dry weigh t pr eser.ted (1':3.ble

2.1+.1.1) is excluding roots. t)ercenta[~eof'cssimLltes out of to t.xl

dry weight diverted to tubers were Lowe st .in tr-eatment S11. It appe ar-s

t.ha t 1D given 'After 3'7duy s uf emer-gence (treJtmer;ts: 1SL"nd SL2) did

not ~ffect the assimilate distributio~ as indicated by the percentage

of tubers out of total dry vre i.ght.

'r",ble2.4.1.1. The effects of photoperiod on some morphologic:J.l

characters of notato.

Ratio ofSpecific leaf 1eaf area leaf to stem Stem drl1 Total dryar-e a dm2 g-1 dm2plant..,,1(w t basis) w t g plant w t g plant-1

Days afteremergence

66 49 '-,oo 49 ,-,-00 49 66 49 66

Treatment

1D 2.18 1.91 40.3 63.2 1.45 1.26 12.85 26.19 32.6 61.4

1S 3.52 3.03 5'1.3 42.G 1.68 1.90 9.64 7.50 34.9 53.3

1S1 2.94 2.51 1+4.2 38.3 1.66 1.86 8.92 8.35 37.5 74.1

SD 3.6LI- - 3.71 1+8.0 41.8 2.10 2.19 6.30 5.17 43.4 52.0

S11 2.39 2.04 45.8 55.'? 1.34 1.41 14.68 19.60 39.2 63.7

S12 3.41 2.48 40.8 29.5 1.61 2.51 '1.94 4.74 43.6 73.7

SED 0.456 0.179 ?48. 4.85 0.211 0.149 1.794 1.451 6.89 7.42

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1'.',

crable 2.h.1.2. The effects of photoperiod on dry we igh t 0" •J. •

(a) tuber, -1 (b) stolon, ' ;.. t--1cr' nlaEt g0 .t. , .~)..Lctn \J ,

and CC) percentage of tubers out of total dry \'le igh t.

:l b c-Days after49 66 4·9

,..,/+9 66emergence bo

Treatment

LD 0 0 1.16 2.15

L3 8.0 31.0 1.13 O.,?O 22.9 58.0

LSL 12.0 49.6 1.81 0.'71 29.2 67.3

SD 23.2 35.0 0.68 0.56 53.5 64.8

SL1 3.3 14.8 1.65 1.75 8.0 21.5

SL2 22.8 56.1 0.62 0.63 46.6 76.5

SED '7.86 7.78 0.437 0.546 15.85 9.23

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37 0 LD 375 o 8D0 L8 o 8L1 flJ

r--j 6. L8L 6.8L2 /I 300 /+' /~

J:3tilr--j

,/ 8EDsP<,/

El 225 qrCJ

" ,..c:+' I I,bO~(J) 150r--j I ~ I~0 (5'r--j0+'CIl 75 75I::''';

til~

0 05 15 25 35 45 55 65 5 15 25 35 45 55 65

375 0 LD 375 0 8Dr--j 0 L8 0 8L1t 6. L8L 6. 8L2+' 3001::' 300

~tilr--jP< /El

/CJ

" 225 225 /..c:+'bO /q,Qj

0I::0 150 150 / SEDsr--j0

/+' \.CIl 'A 1/..c: \CJ

~1::. 75 -b 75 6. ...til1-0~

0 05 15 25 35 45 55 65 5 15 25 35 45 55 65

Days after emergenceFigure 2.4.1.1 The effects of photoperiod on total length of the

main and branch stolons.

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~I

+-' 45 0 LD 45 0 8Dq(Ij 0 L8 0 8L1rl0..til f::,. L8L 6. 8L2q 90 36 36rl /0+-' ~ /til

q I \ /'r! 27 \ 27 8EDs(Ij / /s I /'1-i I I0

/ /1-.

10---6QJ

~18 18

I,¢~ I!:sIr/J

9 9 0'

o5 15 25 35 45 55 65 5 15 25 35 45 55 65

90 0 LD 90 0 8D 0~I 0 0 8L1 /L8+-' /q 6. L8L Lt. 6. 8L2(Ij /'_' 72 720.. /tilq ¢0rl /0+-' 54 54til /~ / 8EDsoq

/til1-..0 36 36 I /'1-i s:0 1,/1-. "1:1QJ.0 Cf§ 18 18~ 1/

0 05 15 25 35 )-15 55 65 5 15 25 35 45 55 65

Days after emergence

Figure 2.4.1.2 The effect of photoperiod on the number ofmain and branch stolons.

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rlI

.p 420 o LD@rl 0 LSp.

8350. ~ LSL..

D:l<:r:

~ 280

_ f:::,

10

O~ ~ __~ __,---,---,15 25 35 45 55 65

25 0 LD

o LS

~ LSL20rlI

.p~Qj

15rlc,..

~ !~

10J0

'-t(l).o~

5J:<!;

I

I0.,'15

b '. _.8- ___

./ ---./ ---'0_Rf-I---.----r-------T .,-

~3~5 (~~~~25Days after emergence

o SD

o SLl

~ SL2

/I //

II/

/I/

95/

I,

.I

/

/

I SEDs

.0

/

'iO /r /

O_II~-~15 25 35 45 55 65

~/~-," ~

O+..J..L___-,---.15 25 35 45 55 65

Figure 2.4.1.3 The effects of photop~riod on number and total lengthof the axillary br~nches (AB).

o SD I ISEDs

.8o SLl ~/

~/

/

~ SL2 8

210,

140_

25

20

15

10- f:::,

5

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~ 1800 0 LD1

+,~~1500ElC)

[/)"1200(I):>(\j(I)~G-; 900o..c::+>b.O 500~(I)~r-I(\j+> 30008

180 o 8Do 8L1A 8L2

18EDs

o L8A L8L 1500.

1200_

500

300

o B I~~'15 25 35 45 55 5515 25 35 45 55 55

210_ 0 LD 210 0 8D0 L8 0 8L1

175 A L8L 115 A 8L2 I8EDs~ YJ1 140_ 140 /+>~

/m~P., /" 105 105 I /[/)

(I):> /(\j(I)

_0~ 70._ 10G-; /0 I/~ A(I) ,35 f(f.0 __ -e- -;-,~ -B 35 /9~ .- -e-:. - /.- [:"

I/ ~--=-=--a•0 0-,15 25 35 45 55 55 15 25 35 45 55 55

Days after emergenceFigure 2.4.1. 4 The effects of photoperiod on number and total length

of the leaves contributed by axillary branches.

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0 L8 0 8Db. L8L 0 8Ll

b. 8L2(J) 100 fT- 0- -0- - - -i!l 100H(IJ I,0;:l

I+"ro 80 / 80(IJ

~ (/J0G-i

ID re- -0-6- - - -e(J) 60 I0 60 I,.q+" I(J) I+" Is:: Ittl 40 I 40rlP.,

6 6G-i0 I(IJ Ib.O 20 I 20ttl I+"s:: I(IJ ICJH I(IJP.. 0 0

20 29 38 47 56 65 20 29 38 47 56 65

0 L8 0 8Db. L8L

7\0 8Ll

30 30b. 8L2

I \24 \ 24lA _

I -~"'-A 8EDs,_ r \I

+" 18 I \ 18~ ~rl \,P., p"H(IJ 12 12.0§ /s::H(IJ 6 6.0;:lE-t

20 29 38·· 47 56 65 20 29 38 47Days after emergence

Figure 2.4.1.5 The effects of photoperiod on number of tubers andpercentage of plants those formed tubers.

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1"./

2.4.2 .sxperimcl!.t Gl~2

In general there was no Lr; teraction between phc toper-Lod and

Lr-r-adi ance levels except in f'ew par-amete r s of the pLant gr-owth , thus

the results in gener-al, ur-e presented c..s the effect of photoperiod and

irradi3l1ce levels.

2.l~.2.1 Stolon

Number of mair: stolons (average) went up to 19 pla.nt-1 and was

not affected by any of the tr-ea tment s ; irradiance did not affect their

total length ei there After tuber initiation growth of main stolons in

SD was negligible wh i l,e in LD they continued grO'.-lir-g(Fig.2.4.2.1).

Branch stolons started coming little before tuber initiation. HI in-

creased the total number of branch stolons as well as their total

length in LD whiLe 11 Lncr-e ased number .u.d their total Leng th in SD.

T':J.;:ing SD arid LD together I HI .incr-e ase d slightly number of branch

stolons as weLL as their tobl length (Fic;s.2.4.2.2 arid 2.4.2.3).

LD increased number of branch stolons and their tota.l length \Vas also

more (Figs.2.4.2.2. and 2.4.2.3). Specific weight for m3.in stolons

went up to 2.4mg cm-1 of its length'lnd of branch to 2.03 and was not

nffected by ,my of the treatments. Data for total stolon weight is

presented in Fig. 2.4.2.4 WId was much higher ih LD, 3fter tuber

ini tintion. St.o Lo» we ight \'/:3.S also sligh tly h igher in HI as compare

to LI.

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20

2.4.2.2 Stem

LI stimulated extension of the main stem (Fig.2.4.2.6) and little

after tuber initiation (TI), effect was more in"LD as SD almost stopped

growing. Rate of appearing axillary branches (AB) in SD was slightly

lower (Fig.2.4.2.8) and difference increased after TI. AB present in

SD almost stopped growing after TI, thus at the time of final harvesting

their average length was 11.52cm while in LD it was 21.25cm. Total

length of AB; wh ich is product of number of AB present and their average

length, is presented in Figure 2.4.2.7 and was much higher in LD.

Effect of irradiance on number of AB vias variable; more AB were present

in HI before TI, after that rate of appearance was h;gherin LI. Their

total length was higher in LI (Fig.2.4.2.7). Data for total stem

weight is presented in Figure 2.4.2.5. 51 days after emergence SD

stopped diverting assimilates to the stems as their weight was almost

constant, rate of stem growth in LD was very high after 41 days of

emergence. Irradiance did not affect the total stem weight at all

(Fig.2.4.2.5)

2.4.2.3 Leaf

Photoperiod did not affect the internode length and length of the

main stem and total length of AB were higher in LD (tigs.2.4.2.6 and

2.4.2.7), thus no. of leaves present on the main as well as AB were

higher in LD (Figure 2.4.2.9). Increa.se in stem length (Figs.2.4.2.6and 2.4.2.7) due to L1 was prirnarly due to extension of internode

length, thus there was no difference in number of leaves due to irrad-

iance. Leaf size in general was not affected by any of the treatments

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and aver-age Le rf s i ze for the Leaves on main stem 2went up to 2.05dm •

Total leaf area was much h i.gher 2.1: I.D espe ci alLy after '1'1, while irrad-

iance levels did not .if f'ec t it (Figure 2.4.2.10). Le ave s we.i.gh t did

not increase in proportional to leaf area thus resulted in difference

in specific Le af i).re.·~(FiC.2.4.2.11). Eoth LD ;md FI de cr ease d spec-

ific Le af are:~, therefore they .ir.cz-ease d , either leaf thickness or

celluLrr dens.i ty or stored car-bohydrv.t.e s or cornbin.it i or, of these factors.

Ef'f'e c t of irr;ldi.:.mce :':':ld photoperiod 0;1 LetlssimiL.ltion r ute LiAR) is

givenb Figure 2.4.2.12; wh i.ch ahows that leaves were more efficient

in producing dry mat ter- ill tr-e.strnent lII. This may be explai ned by the

fact that more pho tons were available per uni t LA. Lower NAR in LD

could be a t tr-Lbu ted to the mutuul eh.rd i.ng of the Le ave s due to higher

LA. LD increased leaf to stem r:ltio in favour of stem especially :lfter

'1'1 (Fig.2.4.2.13). Al though irr:tdiance did not affect the total stem

weight (Fig.2.h.2.5) but leaf weight 'das higher in HI, wh i.ch resulted

higher leaf to stem ratio (Fig.2.4.2.13).

2.4.2.4 Tuber

Tuber .in.i tic, t i or; ('1'1) was exam i.ned by removing ve rmicul i te , at

intervals of 3 to 5 duys , Lr-r-ad iance did :1Ot af'f'e c t TI in combination

'>'Iith SD, where nLan ts ini t i a ted tuberS:lfter 32 days from emergence,

but El e nh.uiced TI in LD where it \'Jas 38 d.iys aftor eme r gence , Some

of the plants unde r tr-ea tmen t comb i.nat ion of LD and L1 intitiated tubers

after 41 d37S of emer-gence but :.t:'..l·)=.:cnts did not h.ive tubers even when

h.:c,rvested, 51 days after emer-gcnce I ',hen 3 out of 4 pl'Jlts had tubers.

In g. erloral tot aL number ,'~.'"'v.re Ll, :_~C" Ch . t 1.. •• h . S_. d - -._ _ ~._, L_...Clr ':lClgfl' 'das mUCI.!. 11lg er In D

(Fir. 2 4. 2 111), U. • I. . -r _ El i'lcre:"sed tuber number in LD Gut decr-e.xsed in 3D.

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Tuber we igh t was Lncreased under both photoperiods by HI but effect was

more evident in LD.

2.~.2.5 Tot:~l dry m:,tter :Jccumul'Jtioll

Roots could not be sepa.r;ltedfrom the peat so t.otal, dry we igh t

(TD'lnpresented in F'igur-e 2.4.2.15 is without roots. LD increased TDW,

which was affected in two ways: due to higher leuf area, they inter-

c~pted more photons and further they had more time for photosynthesis.

Later in the season leaves started coming off the stem and were not

included in TO',I. HI also Lr.cr-e ased TD'l!pr i.mari.Ly due to more number

of photons avai.Lub.le , which increased I!AR also (Fj_g.2.L~.2.12).

Little after tuber initiation there was no competition for assimi-

lates between tubers and huulms in SD as indicated by the evidence that

stolons stems and leaves, almost stopped imparting assimilates

(Figs.2.4.2.4; 2.4.2.5; 2.4.2.10; 2.4.2.11), which resulted in to a

higher percentage of tubers out of TD'I'!(Fi;:-;.2.4.2.16).HI also in-

creased the proportion of tubers out of TD'\·!(Fig.2.4.2.16). In SD

effect was one way, as there wus enough induction so percentage increase

in tuber weight was due to higher ouan t i ty of assimilates available.

In Lu it might have affected in two ways: one by stimulatir.g tuber-

ization and another by rtla':d!lg more aas imi.Latee , ava.iLabl.e,

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240 SEDs0 SD

o LD/0

200 //

;- /- /- -0160 I,_I /+'

s.:::

/'1lrlP.

S 120 I /o

"..s::tos.:::Q)rl 80s.:::0rl0+'Ul

40

o ~------.-------.-------r------'I-----------'6525 45 8035 55

Days after emergence

Figure-2.4.2.l The effects of photoperiod on total length of

the main stolons.

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90 Irradiance 90 Photoperiod r0 HI 0 SD0 11 0 1D /

75 75 /r--II

/+> 601/ I SEDsi:l 60til

r--IP<

/I-. 45 45Q) r: ---0.0

~i:l /i:l 30 30 /0 Ir--I

/0+>

~Cl) 15 15 ""0 025 36 47 58 69 80 25 36 47 58 69 80

Figure 2.4.2.3 The effects of photoperiod and irradiance on numberof branch stolons.

Irradiance Photoperiod550 550 70r--I

/I Cl+> 440 440i:l /tilr--IP<fq /u 330.. 330

I / I SEDs~+>~§3 /r--I 220 220 ?-- --0i:l0

/r--I0+> 1,10 ./Cl) 110

_A,/

0 025 36 47 58 69 80 25 36 47 58 69 80

Days after emergence

Figure 2.4.2.2 The effects of photoperiod and irradiance on totallength of branch stolons.

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Irradiance Photoperiod1.8 1.8

HI SD P,..... II LI Ld+> 1.50 1.50 /§ /..;~ 1.20 1.20 /bO

" / I SEDs+>'tb ·90 .90 I II'r-! }l IQ)~ / ?---ci1:: .60 / .60'tj

q0..; ·300 .39+>Cl)

0 025 36 47 58 69 80 25 36 47 58 69 80

Figure 2.4.2.4 The effects of photoperiod and irradiance ontotal stolon dry weight.

35 Irradiance 35 Photoperiod,_ fJI

28 0 HI 0 SD /+> 28q /Cl! 0 LI 0 LD'it /bO 21 1/ I SEDs"

21}+>

'tb'r-! /Q)~ 14 14 /:s '/

l1'~ /Q) /+>U'J '7 7 0/ ~

...121 0

0 025 36 47 58 69 80 25 36 47 58 69 80

Days after emergence

Figure 2.4.2.5 The effects of photoperiod and irradiance ontotal stem dry weight.

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50 Irradiance SOl Photoperiod I SEDso HI t 0 SD BI

I o LD 1//

40 o LI .B 40J../ J21

/a 30 ,0 3°1

/I/tJ /

// r 0

" 0'/

I~

..c::of.:> ./IlO /I:l 20 20<Ii rt.-;

av~~:of.:>

Ul 10 10I:l ;:J~r'rl

~

0 0-,---, ---,10 24 38 52 66 80 10 24 38 52 66 80

Figure 2.4.2.6 The effects of photoperiod and irradiance on maJ.n stem length

.560-'.l Irradianceo HIo LI

ILl////

.0./

./)2r

/j

-::;::;-

O-..;--,------.·.r----,--- I

J(; 4'1 :3H Cl') f5()2')Days after emergence

560_\ Photoperiod Po SD I

. /81 0 LD I44 .~ I\ II1 JZl336

1 •

I I/2 2 (~'1 {Zf

I II112--1 'I

I .21 ~

I// »-cr ./~-

O'-rG=~-r---'----r--25 JG 41 58 69 80

ISEDs

·Figure 2.4.2.7 The effects of photoperiod and irradiance on totallength of the axillary branches (AB),

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Irradiance30

25,_I

-f-) 20$::1alr-i~"

15~c.....0 10!-.(J).oS 5~

025 36 47

Figure 2.4.2.8the number

58 69 80

Photoperiod30

o SDo LD

25 I

//

I / I SEDs,B

."..

20

15

10

5

o25 36 47 58 80

The effects of photoperiod and irradiance onof axillary branches (AB).

ft- -I:]-:20 (a) /' 225 (b) 110 SD f SEDs 0 SD /

,_LD /I 18 0 LD / 180 0 /-f-)

§ ,I " SEDsr-i~ 16" J /!-. 135(J) /.oS 14 /$::1

c..... 90 ;Ial(J) 12hi

10 45

8 025 36 47 58 69 80 25 36 47 58 69 80

Days after emergence

Figure 2.4.2.9 The effects of photoperiod on number of leavescoming from the main stem (a) and the axillary branches (b).

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100 Irradiance 100 Photoperiod SEDs,_I 0 HI 0 SD 11~ /J::: 0 LI 0 LDtil 80r-i 80 /Pt

C\J /~ /

" 60 ~til 60 ./OJ~ ./til

~~ 40til 40OJI-=l

20 20

36 47 58 69 80 36 47 58 80O'_--~----.---~----~---,25 o

25Figure 2.4.2.10 The effects of photoperiod and irrandiance on

total leaf area.

HIPhotoperiodIrradiance

o o SDLD,.._4.1

I

o LI4.1

2·90'

_-El....<,

I SEDs

2.90<,TI-_

25 36 47 802.5014---~----.---.----r--~

2558 69 36 47 58 69 80Days after emergence

Figure 2.4.2.11 The effects of photoperiod and irradiance onspecific leaf area (SPLA).

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Photoperiod Irradiance

2.20'r-!+>til!-t

~ 1.9+>til

0+>~tilQ)H

2.2

1.9'0-_- --_

~ ...........................

"El'\lSI\\\

o SDo 1D

HIoo 11

1.3\b----El

1.0~----~---'----'---~----~2525 36 47 69 80 36 47 58 69 80

Figure 2.4.2.13 The effects of photoperiod and irradiance onleaf to stem ratio (dry weight basis).

Photoperiod Irradiance4.00

3.40

2.80o SD

1Do HI

o 11C\JIEl

~ 2.20 /1\/ \

\'\

o

1.60

1.00+-----,---"'"1-'-----'1"'----r __-----w

2536 47 58 69 47 58 69 808025Days after emergence

The effects of photoperiod and irradiance on netFigure 2.4.2.12assimilation rate (NAR).

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Se.:)

rlI t: t:.+>s::tilrlP.

bO

" 33 I+>..c:eo

I'r!QJ SEDs~~ 22H'Cl

HQJ

~I:;

1·1

50_

~orlI

+>s::til·rl 30_p.

" I IH IQJ SEDs.0

3s:: 20.HQJ.0;:jI:;

10_

Days after emergence

Figure 2.4.2.14 The effects of photoperiod and irradiance ontuber numbers and tuber dry weight.Key: 0., SD; 0 ,LD; closed symbols, HI; open symbols, LI

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Irradiance Photoperiod

8 0 HI 0 SD800 LI 0 LD

,_ 64 P 64I /oj.) /§7i. 48 / 48/C) ,;i I';3:.'"

SEDs~ 32 32

16

80o

25 36 58 69Figure 2.4.2.15

16

o

total dry weight (TDW).The effects of photoperiod and irradiance on

80 Irradiance 80 Photoperiod0 HI 0 SD0 LI 0 LD

64 64 I SEDs

';3:.

~ 48 48~0oj.) -::'l 32 ..... ......0 32'"~ 1Y(j)

.o::'l 16 16.~

~ ,..o-----[]......-

,/

0 025 36 47 58 69 80 25 36 47 58 69 80

Days after emergence

Figure 2.4.2.16 The effects of photoperiod and irradiance onpercentages of tubers out of TDW.

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2.4.3 Experiment GH1

Due to fewer number of replications there was more variation in

the data.

2.4.3.1 Stolon

I:umber of main as well as branch stolons and their total length

was higher In control (Table 2.'+.3.1).

2.4.3.2 Stem

Length of the main stem is presented in Figure 2.4.3.1. All the

three shades used at i.mul.ated stem extension and difference was visible

even two days after moving them under the sh:tdes. Control had higher

number of axillary br~1ches ~d their total length was also more

(Table 2.4.3.1), but difference \V3.S not si.gni f i can t due to variation

in the data. Although plants were taller under shades but total stem

we i ght was higher in control (Tc,ble 2.4.3.1).

2.4.3.3 Leaf

Increase in Lengtlt of the rnoin stem by shades \-lelS purely due to

ex teris i on of in ternode, ,is there "Jas no difference in total number of

leaves (yellow or deud + green) recovered from the muin stem (Table

2.4.3.1). No. of Leavea present on axi Ll.ar-y br::mches were higher in

control and leCl.veswere bigger in size in control and their total leafarea was also more (Trible 2.4.3.1) Lncr-ease in Leaf we igh t was not in

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proportion to the leaf area, resulting, differential values for specific

leaf area. All shading increased specific Le af area (Table 2.4.3.1).

Leaf appeared slightly thicker in control but were not measured.

2.4.3.4 Tuber

Plants under muslin sh.ide did not form tubers at a.l L, while in

red shade there was only 1 tuber plunt-1 as compared to control where

there were 12.7 tubers plant-1, tubers dry weight for red shade vias

negligible (2.7mg pLmt-1), while in control and blue shade s it was

1.42 and o.4g pL-mt-1 respectively (T·,ble 2.1+.3.1).

Tot·od dry v/eight

Control had higher total dry weight (T:;ble 2.4.3.1) and quan t i ty

reduced by sh.ades was a.Lrnos t propor tionuI to the number of photons

t r anemi tted through the respective shades. Percen tage aas imi.Late s

directed to the tubers were eoua.l in CO!l trol .md blue sh.ide a,

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c...:.,;

'I'abl,e2.4.3.1. The effects of different type of ehad i r.g on morphologicoU

chur-acter-s .md tubers, Lt~) days after emergence.

Treatment cOElrol mus l i r: I\Qd Blue SiD

r:o. of axi Ll.ar-ybranches (AB~Pl -1 1+. '/5 1.0 1.33 0.33 1.728Total length ofAB, -1 22.7 6.lf 5 P 2.2 '7.73cm pI .,_)

\0. of greer. leavesmain 1-1 13.25 11.6'7 11.67 9.33 1.214on stem, p

I'~o.of ye Ll.ow or deGdleGves coming frommain stem,pl-1 0.50 2.6'7 2.6'7 4.00 0.894Average siL;e of leavescoming from ma.i n stern(dm2) 1.03 0.61 o J,~ 0.56 0.096• T i

S'Jecific le::!f"re" dm2,.-1 2.66 3.94 4.82 5.02 0.134::<. c•.r.. , d G

r;o. of main stolons, pI-1 15.3 10."/ 10.0 9.0 2.96Length due to mai::lstolorls,cm pl -1 57.1 24.6 1?9 21.1 1'7.48No. of br-anch stolons,1-1 r?3 3.3 0.0 O.? 2.91')

Length due to b~,:.~nch9.3 1.6 0.0 0.6 3.31stolons, cm p1-

'l'uber b 1 -1 12.'1' 0.0 1.0 4.0 3.35num ers,o __

Tuber weigh t, -1 1.42 0.0 0.002'7 0.40 0.304dry g DlTotal dry weight, g pl -1 10.57 3.91 3.16 2.9'7 1.035Tot:J.l -1 3.18 1.62 1.63stem \'.eight,gul 1.32 0.39'rota.lstolon we i gn t ,g :91-1 0.33" J.O(' 0.05 0.03 0.11Percel:tage of tubers outof TD'd 13.4 0.0 0.1 13.3

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48ao~

.r::+'eo>::: 32OJrl

SOJ+'til

16

80 SEDsBlue shadeRed shade

Muslin shad

fJ/

/

0-4-_- ,-- ~~---I-------~-- ---T---·----~---- r~~-----·--r--·~---~---T---'5 11 11 23 29 35 41 41

Days after emergence

Figure 2.4.3.1 The effects of shading on maln stem length.

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2.5 DISCUSSION

There is ger:eral agreement between experiments: GIll and GR2 that

SD stimulated tuber:bzation (even there were some s ess iLe tubers) arid

decreased haulm growth after tuber initiation. In Experiment GR2

where growth analysis was done regularly, after 41 daya of emergence

there was no increase in len~th of stolons or stems ~:d ~fter 51 days

of emer-gence there vias or.Ly r.egligible growth of stolons, stems and

Leave s Cin term of weight) so iumost etll the assimilates produced, were

be i.ng directed to the t.uber-s , It may be easy to explain this after

understanding the theory of tuberization.

I t has already been mentioned (Chapter 2.1.1) thu t Imler levels of

gi bberellins and higher levels of cytokinins were de tected in SD as

compare to LD. Further exogenous u.pplic::J.tiol1 of gibberellins have

been found to inhibit t.uber i za t i.on (Lovell and Booth, 1967; Hammes and

Beyers, 1973; l-lenze L, 1980), on the other hand Pal.me rv und Smith (1969,

19770) demonstru.ted the requirement for cytokinins in tuberization of

excised stolons of Solanum t.uoer-osum .in vitro; similu.r results were

obt aLned by Haux and Langile (19'('8), using Ziatil: riboside instead of

kinetin. V/areing and .Ienn i.nga (1980) found that er:dogenous Abscisic

iicid (ABA) sUP:9lied by the le:J.ves \'l:J.Snecessdry for tuber development

in Solanum and i gena , but there \.j:_;._s LO s i gn i f i can t difference in ABA

contel1t in diffusate~ from induced to non-induced leaves. Further

more a non·induced le:J.f could induce tuaerization when grafted on to

an induced stem cuttings ('.'iareing and Jennings, loc. cit.). Supply of

exogenous cy tok in ins plus AJA did not cauae tuber f'ormut i.on in one-node

cuttings f'r-om.a non-induced ~)lCl.ntof Solullum :mdic,cu.l (ilil.reing and

Jennings, 1980). They argued that the clones of SoLmur:1 andigen/i., they

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used have &11 obligate short-day requirement where as many cul, ti var-s of

Solanum tuberosum have no ilbsolute requirement for SDi but give a

quantitative SD response. Thus it might be possible that a second

factor \13.Spresent L1 Solanum tuberosum .indepe nderrt.Ly of the photo-

periodic pre-treatment. Thus it may now be concluded thc t the tuber-

ization in potato is promoted with increase in ratio of cytokinins

and unknown tube r-Laa t ion s t iuul.us (UT.')) to gi.bber-eLl ina , Production

of the UTShas been suggested to be promoted by SD (';Jareing, 1982).

Tuberiz3.tion involves cell divisions in the sub-apical region of

the stolon apex (Booth, 1963; Cutter, 1978). Cytokinins, have been

suggested to Lnh i bi t .rp'i cuL meristem and promote cell division in the

sub.apical region (Palmer and Smith, 1970); thus with increase in levels

of UTS and cytokinins, tuber ini t i at i or; occurred and w i.th a further

increase in number of short days, the ratio of cytokinins and UTS to

gibberellins increased. This further stimulated cell division in the

SUb-apical region of the stolons and theJ._pical meristem might have

been inhibited to the extent that stem and stolons stopped /Srowing

and aJ.I assimilates produced by the leaves were directed to the tubers.

In Experiment GR1, after 35 days of emergence treatment LS had

3.64 tubers per pl ant and 85. Z':' of the pl ants had formed tubers, while

in treatment SL1, there were only 1.5 tubers per pl an t and 66.7% of the

plants h3.d tubers; but both of them had equCtl number of short days by

that t irne , with the di t'f'e re nce that LS '.'las given short day after 17

days of emergence and SL1 straight af te r emergence. Furthermore

number of axi Ll.ar-y br anche s , their total length, no. of leaves on

Qxillary br anche s and Le.ig th of stolon, c.ll were higher in S11 as

compare to tr-ea tmerrt LS. In:ln ano the r treatmen t , 1SL, which had 20

short day cycles but af' te r , 17 days of emer-gence ; tuber we i.ght after

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49' .ind 66 days of emergence \Va.s23.2 and 35.00g 91:'l,t -1 whiLe in SL,

4 8 -1 .it was 3.3 and 1 • g p.lan t r-espe c ti ve.ly , Stem we igh t which is in-

creo.sed by non-inducing conditions was also higher in SL1 thM LSL.

Specific leaf .J.rea, Vl':{S Lower .in 31,1. fes we have seen ear-Li.er that

effect of photoperiod is ou.m t i t a t i ve , thus difference of only 3 short

day cycles cannot be the only reo.SOD, thus there must be some other

factor affecting it.

l!urtJ' and Sahu ("ococ::), . L... 0<-,., I -' (./ , found thut 20 SD cycles Given immediately

after emergence or 20 d:~s o.fter emergence, failed to initiate tubers,

while 15 SD cycles were enough when given 40 or 60 d.iya after emergence,

wher-e plants were ke p t .ir: continuous light and harves ted after 90 days

of emergence. In ,:'1.11 expe r imen t where pl an ts in .rddi tion to '? old

leaves had: 3 young (less thai: 3cm) t.er-m.i na.l Leave s or one young

termir:al leaf or \·,ithou t·,ny terminal Leaf , Tuber wei ght was highest

where there wus no young term.~Yl:"'.lLerf ....nd Lowes t wher-e there wer-e

three young term.' n'11 Leave s (Hummes .md Beyers, 1973).

Thus it follows that the ratio of production of cytokinins and

UTS to gibberellins is more in old leaves compd,red to younger ones

'-'.nd obviously ratio of old to young Le.ive s Lncr-ease s \-.rith age of the

plan t ,

In case 0: tr-eatrae n t LD: ill Expe r imen t GR1, there W'.lS no tuber-

iZ,Dtion until 66 duys:·.fter emergence whei: this expe r imen twas term-

in:~ted, whiLe in Exoer imen t GR2 under h igho r irr;~di'Ulce' level (l~O 0

U:'H-2 -1..c,:,: S , -1still lower th~n GH1), some tubers were recorded (10g plant ) •

This clearly shows that tuberization is f~voured by lower temperature

as in Experiment GR2, tempe rutur-e ,-J,),S i5°C during day .md night while

it. ExperimentGR1 temper-at ur-e \1;').S

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Resul ts are ill genera]_':lgreemen t '.vith pri vious workers (Gregory,

1956 i Bor-a....1J. and !'!l'lthor'_,,)e, 1062' Sl. ter 1q68· Q.,}, ., et .,1 197tc•~ ..I , --'."_ , " ,U:"'Ul.'_.\. ':......-L. , r,

l'~enzel, 1980).

It t lower irr:l.dia.nce level i:1 expe r-Lmen t Gl~2 .ir; combi n-it ion VJith

-1 QLD, tuber weigh t ·..ns O.5.~ ClLl'1t af t er u1 days of emer gence but ,··~t

hieher irr:ldiar;ce level i» combinat icu "lith LD tuber weigh t vn.s 14.5g

pl,::l.l1t-1• SimiL:crJ.y in expe r imen.t GH1 pl:.cr:ts under musl i,» ahade did

not 'tube r i.ze whi.I.e p.l.anta grov/il:g in n...t.ur-al glass house .rt t ained

1..422: of tuoe r s per ?lant (there v;;_~sno difference in ape c tr-al, photon

distribution). Similar resul ts in Growth rooms were obtaine d by

Borah, (1959) in vari e t.y Arrun Pilot (irr:.Hii:.Ulce levels wer-e 61+ and

128 co.I cm-2 day -1). Either induction i.ncrease d I,!i th Lnc re ase in

irradicmce levels, or D.l ter-nati vely, tuber induc t i.on was a.l r-eady there

and the add i t i onul, aas i.miLat es ava i.LabLe due to higher number of photons,

were diverted to the tubers. F~vouring the first~y?othesis, analysis

of endoge nous ~ormones .in SOl:J.llUr.1:.u,dir;en:J. showed t.h.rt Low light

intensi t:y Lncr-eased levels of .ic.i.d i.c Gibberellins in Leave s of short

day pLmts ('~;oolley and 'dare i:1g , 1972). 'I'lrus under higher light intens-

ity ratio of cytokinins a~d UTS to gibberellins might have been higher,

""hich resulted hit:;her tuber \'Jei,;ht.

'rot::li dry mat ter Droduced ',idS reduced oy decrease i1: irradia.n.ce as

earlier reported by (Pohj;"hk,.~llio, 1951 ; i3odl:.\I~r"der, 1963; Sale, 1973;

BorQh,1959). NAR'lias low in LD due to mutuQl shuding a~d high in HI due

to higher ~umberof photons available for photosynthesis. Specific leaf

o.reo. increased with decreo.se ill irro.dicu,ce levels; e.g. l'~x8eriment GR2

and GH1, o.nd vias hig:,er ii, S">as com::>lre to LD 0.5 fouEd, oy Dorah (1959).

R,,,tio of le2ves to stem (o~, \'!eiG~;_tb:.lsis) decreased i,: LD :lfter tuber

initiatio!1, due to co:. tinuous f.;rm·,tL of the stem.

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jU

cre~lse in Lr.te rnode ler,gL1 (0.'::. comn.u-e t r e itmen t a 701,).] ':r in Exper-

iment GE2 ar.d rnusl in v.nd control in Exoe r imeut Gi~1). Iccre:Jse in stem

length of po tut o due to de cr-e.xse Hi r(d,Lt'cOl' :kS',,:LSO be en r-enor te d

es.r Li.e r- (Bore11, 1959; BodLender, 1963) .ir.d m.iy be reL,ted to higher

UJ;:ount of cibberel1ins "reduced under Lowe r irrHl.:i:,flce ::'evels Clolley

de cr-erse ill phy tochromo .sL.,~e. For exunol e , iu crc',trnent bl ue and

m~slin shide ir. Expe r imer.t 3:11, to t.il r adiut i.on '11<,,8 the sume , but plants

were t.rl.Le r unde r blue ch:«lc where Ohy Lo ch rome s ta te ',-1,,),3 Lowe r than

Ul'.der muslin. Stirr.uL"tory r-e aponee CLell::r,th) of flr red ( >'700nm) and

Lnh i bitory of red Ugh t «,?OOr:m) is es tvbl i shed i~'l v.crious crops

(Imr:oft et J.l., 19'(9; Jtcques, 1)68; Satter .ind '.,'ether'1.J.l, 1968).

Effect of sDectr~,l nhoton di e tr-i bu t ion 0;: t.uber i z.at i on is not QuiteL '. ;.

cle·'.;r from this exoe r irne n t :J.S to ta.; r,di :;,t i or, w.isicon Iounded vii th light

nU::;,_lity. Howe ver blue li.::::llt :lQOearsto howe SOI:1ea t i.mu.la tor-y effect on

tuoeriz",tio{, ,1S per-cen t"se of .:,ssimiL,tes diverted ;;0 tubers were equul.

in pl!Ul ts grO'.-JEunde r blue sh.ide s .:Jl1J n:.l tur:.l C;l',;,ss ;1ouse while irrad-

ie.nce 1'lCeS much less ur:.der blue sh:)de. Fur~her there '.-:3,S no tuberization

in plunts growr. under mu.sli~ or red shade.

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3. CHEMICAL CONTROL OF GROWTH

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j1

3.1

c:r:Ol'!!l tubers m·.·.]r-espond i.Jy CC'S.il~G to (JU1i;:·.md 'OJ grmvir:c out s tolor.s

(E:-.mmes .md l.e L, 1975; i'~eLzel, 1980; l.ur t i -md B'lr~erjee, 1976b;

1962; Tizio, 1?64; Dyson, 1965).

The .i nh i o i tory effect of G;, OIl t ube r+f'or-mu ti cn h.rs Le d to the

eXoeriment,l ,;!nlic:~tio:; of ;_r,rO'.lthr-e t.vr-de r, t such :'05 ?-chloroethyl tri-

rr.ethy~ ammon ium chloride r ccc i. ',,'her: o lvr.t s we re .eit ae r s)r::yed w i th

of ....er i a.l stems ','.d Le.if .•re. wer-e de cr e.ised (Kr ug , 1961; Jyson, 1965;

GUl:::lsen,,_rld 1I:lrris, 1969, 19'71; Kum.rr .ud 1.!arei"G, 19'/4; JIurti arid

Sah'l, 1975'0), tuber \:ei!,;h t \·Ji.S i!:cre·)seci e sne c icl Ly follmring eee app.l i>-

c:.:tior, (0yso.], 1:;65; Il.urme a ,".•d LeL, 1975; lle nze L, 1980). If eee was

a~)~)lied ar-ound tuber ini t i. t i or: (1'1), E,er, tuber numbe r-s wer-e increased

(Gun·lse::lc. and ll ur-r i s , 196); GifIorLl .u.d j·:OOY·oy, 1967).

The e f f'e c t of ecc is gre:Atcr ur.de r- coud i t ior.s , whi ch s t imu.l a te

,. . .,i.e nze.r (1930) reported greater

incre::1se in tuber weight due to eee.~): .ic, t ion .ir: p1,); t s crowing a t

hiGher temper" tures (32/18 or 32/23°C) com'):lrcd to 10'.:or temDero.ture

'Jhotopcriod:_c ex~)eriment, " unmes.nd Lel (1975)

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reported c;re:\ter .i.ncr-e.ree i?l tuber ','!ei;ht due to eee :p!)lic:.ltioYi in

:oLmts grm'li:JZ .it 18 hour-s cL_y Lengtri corr-iir-e d to 12 hour-s; The effect

of eee was more in L!vour of tu be r-s V::10J: ni troCCYl \-!><] 3)~Jlicd at higher

r a to s (GUJW_sc:j.:~ and Har-r i s , 1969).

The Lnh i c.i tory e f'f'e c t of eee on grO'..'th is tl':.lISi toz-y and de cr-ease s

ste~dily over ~ ~eriod of 42 d~ys ([rue, 1J61). Jhe~ ece wus used at

the rnte s of 5Crnl, '7,)C) or oe r

rr, te of the a tern crm'ith ',\'·!.S Low for .l »e r iod of 30 d.iys I'o.Ll.ow ing eee

a1;)~')lic;:ltio;, and tuber .in i t iut ior; \'!'--~S enh.inced but fiLed tuber weight Vias

not af f'e c te d (Dyson , 1965).

?: -dime th~·l-'l:1inosucc:i.Ll;:liclcid (:3) h:".d simLL,l' effects to those

of eee C;iviqj :.l Ger:er,-,l de cr e.ise i:-, h.rul m grOi,th (D:,'so:: .:'.:ld Humphries,

1966) and tuber numce r cud tube r ero'llth '.-lere Llcrecll3ed :;:;_rticuL:-.rly in

the ')eriod follOlviLg i ts ",~»)licutio,: but fin.ll tuber yield \'I:1S not

,'ffected (Humphr i.ec ~;_rld D~'son, 196';').

A more rece.i t.Ly ay.ithe s i ae d erO':lth rC'::;l1Lltor, ;J)333 (Imperi;.ll

er_emic:;l Lndue t r i.es ) h.is oee , GhO',-Jl". to decr ec se culm lenGth and in-

cre.')_se aeed y i el.d et -,':"'L~. , Up t ake of

~)P333 ','I:lS largely from the soil vi a roots :_,nd .ic t i vi ty, or :~t Le as t the

observed r-esponse "/'':'S m:'.int.:,ined over_, lonL~ period. It wa5 considered

t~(ji3_tthe Gener,l reS::JOlJ8e to :)-0533 ';:~,Ssi:'1il.JT to that of eee and '39

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3.2

effects 0:: Luber y.ioLd s , t. }jj,:-::hcr :)2.'o)ortioll or to t;.»! oLrn t dry mrt te r

in the tubers woul d »r ovi.de the »o ter t:i '11 :or Lncr-e.rse d pr-cduc t ivi ty

vestiGe" ted .ir: si;:;;~_e ;Jlo~s, '..: j t:l se''1uc:1 ti:'_ h.ir-ve s t.s for crop growth

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Y+

The experiments were car r i.ed out en the Universi ty of Kotti ngham

farm, at Sutton Bonington, over the two ye ar-s , 1979 and 1980. In 1979,

soil \-JllS aandy cl.ay Lo.irn ([<'ield 32) '.~Ed Ln 1980, it vuus smdy Lourn

(Field 10).

On receipt, seed (det~il for seed source in Appendices B :md C) of

the vnr-i.ety Pent l.and Cr-own, was examined for di se.rse etc. ln 1979 seed

was infected vii th f.:i1izoctonia sol:mi Kuhu (Black scurf). All severely

affected and deformed tubers were di.acrr-de d and r-emaini ng tubers were

Qrra ..nged 'A~ic:ll' end uppermost in i.l single layer in chi tting trews.

Seed was then stored (Table 3.3.1) untiJ.. pl an td ng and irradiance to

avoid e t ioLat ion of the spr-on t during s tor age WiAS 2.3 ~ O.l~ UEIvl-2 S-1

(warm white fluorescent tubes), on top of the tubers. Rel~tive humidity

Experiment:,l design ilnd 1)ri-{ctici_~.ldetilils

:2reatment de toiLs .ir-e gi ver. in Table 3.3.1. SiEgle app.l.i.cat i.on of

-1PP333 @750iS ~;.i. hu \'JaS gi ven vis i, apr-ay , covering the whole plot

equally i.e. p.lan t.s and bar-e so-il. There uere no cul t i vat i one after

pLurting and weeds wer-e effectively controlled, us ing ~l. mixture of

parG.quot and Li nur-un itt r-ecornmended dose ,J.t about 5~':emergence. Over

the period when stems were emerging, nwnber of stems were counted for

10 plants per plot, OE every other day in the begimiin;g,when stems were

emerging .it fast r ote and on every Ij - 5 duys La te r- all, until no further

increase took p'l ace , Dat;). for r-ainf'cLl , screen temper-uture and soil

temper.:l.ture ,~,t 10<;:r.1dep tli VllS obt.i.ine d f r om :_,ne .u-by meteorological

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a t.at i.on , w i thin one kilometre of the expe r iment.s.l si tee r~dinfcdl

totalled for every five dJjs is presellted in FiGure 3.3.1 and screen

(ma.x. and rain , ) and soil tempe r ature ,~t 10cm depth, oath .rveruge d for

every 5 d~ys ~re prese~ted in Figures: 3.3.2(a,b) ~~d 3.3.3(:..:.,b)

r'espe ct i vely.

Light Ln te r-cept icr. ir. the pho torsyu the t ica.l Ly ac t i.ve r ange

(400 - 700nm) was measured 2'1 situ us i r.g Q,nYltummeters (Lund;.;;.

instrumeEts) w ith remote cos inc corrected sensor heads , SimulLu:eous

r:adings were taken from above and beLow the cr o» cunopy iat r-andoml.y

selected Dositions. Reflect~nce in t~e 400 - 700nm range was considered

to be low (Scott et al., 1968) and relatively constant and consequently

\v:..:.snot included in the r-eudi nga, To decide, the number of readings

per plot; in 19'79, 40 r-ead i.nge per plot were tuke n for few plots of

EXperiment F1 and co-efficient of vari:ltion w~s worked out for all the

40 r'ead.i nge and the r. selecting r-undom.ly 5 .md 10 out of l~O ('rable 3.3.2).

It w,').sdecided to take 10 readings pe r plot. Er; order to mi.nimi se

the influence of solar height, all r-ead inge were taker; between 10.30 -

14.00 hours. Early in the season when ground was not covered completely;

readings were LIken around the p.lan ts and thus per-cent age light inter-

ce:9tion was ~djusted for ground cover, mea.sured using the grid.

The grid consisted of <) r-ectangu.l ar metal, fr:une, w i th strings, at

, 15cm spacings pulled tightly acr-ose tile f'r-ume, Size of the frame was,

150 X 90 sq.cm to cover, 2 ridges. The gr-ound cover ""<s recorded by

~lacing the grid ut two places in Cl plot und looking downw~rd.

Recommended p.lar t protection me.rsur-es were t aken .md crop stayed

hea.l thy dur ing both ye~U's :IS L.r as insect pest .md di se nse s were

concerned.

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3.3.2 GrOlvth an:.l1'Isis

Seven gr-owth :.iI'.:i.lyses were c.u-r i.ed out during bo tn ye.~lrs.

Sampling W&sfre~uerit early in the seaso~, to obt~in ~~ idea of earlier

tuber gr owtn , ~~:.:.tcha.irro l.e co.rs i s ted of, 2 .in 19'79 .ind 4 if: 1980,

adj:lcent p.l.an te .~;nd car-e '.:as taker, th.rt a t I eas t one gu:~rd p.l an t was

left between each si:u;'!Jle. At e ach hir-ve at f'o lixge of the selected

plants Vias cut off at soil leve2_';_wl tne.i unde r-gr-ound p.rr ts of the

plar:ts we re lifted cur-e f'u.lLy , using a fork. All tile tubers, however

small, were recovered. Plants were stored in labelled polythene bags,

~t JOe to. d' t d~ q un l~ lssec e • La ut ternp t w·_,s m~.de to recover r-oo ts umd those

whi.ch we re a t tached to the s tems were removed:,u;d diacnr-ded , Labor-a-

tory procedure for gr-ow th :m·Qysis \'k, co the a.ime ~.lS described in 2.3.1.4,

except that s talons were no t measure d;

K,rly Ln the se·~)sor, the Le.rf .ind s t.er: m.ite r i i.L from the whole

s,'Jnple were used ir: de te rmir.Lng the dry;;, .. t t.er co.rte r.t , 'out as sample

size .iLcreased,

i!lg all tbe s i ze o ;' s·ir.nLe.

·After cornpLe te ae ne ac i.ng , gUtI'd pL.c!:ts on either side of the rOI';

't/ere har-ve s te d w ith :, f'oz-Ic, r-om.rining pl un ts , excluding 0u.~rd rOI·/S

Were har-ve s ted w i.th ,_ tr"ctor ope r-ate d oo ta to digge r (Jonson), and then

picked manua.lLy , 'l'uoer s wer e stored in .labe Ll ed sucks in ~\ cold room

~j_t 4°C, un t iI p.rase d through the riddle. Piddles a·r.ciL(ble for griJ.ding

large qu~ntities of produce were slightly different from the one used

for growth;n.:,lJ'se~. Fecords v/ere r-,.cde Yor the :~ur;.oer ;\1:(1 fresh weight

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3?

Table 3.3.1. EXDeriment~l dettils

19'79

Pre -91:311 t ingstorage.

9"/ days a t 4°C,followed by 13 d~ys ~t120C .md 22 days a t SoC.

S~ed size 105.3 "t 1.31g

Plot size 5 r-ows of 6.56H e ..ch

Tre a. trne nts There we re 5 tre«t-men ts: 1)P333 epr.ryedon; 5th June or 13thJune or 2nd July or16th July illd control(no spr..,y).

h'e t'l' 131""'" h-1... r l aze r uf,g :'.(All given beforePI;~nting) (15:15:15; N:P~5:K20)

Dose of pp333 , , -17'SO'gl.l. nil

1980

90 duys i: t 4°C,followed by 39 dctys at120C .ind 1 eLlY at SoC.

15 th Aor iI

'17.5 "t 1.26g

8 rOIlS of '/.20H each(control)

10 rows of '?20H each(p:J333)

76X36cm (control)G1X20cm (up333)

There were 2 treat-ments: pp333 sprayedon; 23rd Eay :J.ndCall trol (no spriJ.Y).

(15:15:15; I::P,205:K,20), -1750g a , i , ha

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of tubers in each grude .·_ald, aub-zs enoLe of 350g to 4Kg 'lld.S taken,

depending upon the quun ti t.y of tubers in e .ch gr·de, for dry we i.ght ,

T:tble 3.3.2. l<e:m per-cent age Li.gh t .ir: te r-cep t ion be fore a.djusting

for ground COVerl!ld CV:'~,OL 21.6.79.

VARIETY 2entl,ndCrown Record

Renlic.ltionne; of

r e.tdd ngs CV% me un CV%

40 pr + 0.9 1 83.6 + 1.78 2.1300 -

10 84 + 1.3 1.5 83.4 + 3.331 - - 2.77

5 83 + 1. .3 1.5 82 + 3.5 4.3- -

1.lf-2 86.5 : 0.86 0.996

2 10 84.'( -: 1.46 1.73 88.5 +- 0.92 1.06

5+83.3 - 2.6 3.12 86. '76 + 0.89 1.02

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I

I :

------~-----------------~I:~--~I~~·----------------------------~--~--~8I .i.. I

, II f-

It.:11r~

:- 1,-' _...s.'_' _~~f-

rf~..., ------r,-------..L..I1.....j ~~------~----------r_--~~~----~l ~ __~.--~~J -I

I

~-_ .........--1 :>.L-,, __ -13

, t-:J

i

I II (])i:----------------------4--------~I~~§J t-:J

i., I

lI ]: :>.

r-----_,~I1i IIr-------'ll .';:jI Hl --I ~

L..--------r---~--r-_,r__.--_.--_.--_r--_r--~~i 'i

o 0 0 0 0L.I'\ ..::t (Y) C\J o

oco0\rl

'0§

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Ul

ttl>rOLr\

?>1--1Q):>Q)

1--10Ct-i

-oQ)bOcO1--1Q)

+' :>CJ cO0

0\t:-0\rl

eos:1'rl1--1

+' ;:jP. rOQ)Cl)

U0

Q)1--1;:j+'eo cO

;:j 1--1c:r.: Q)p.SQ)

+'

?> 0rl;:j~ s:1

'rlSrOs:1cO

Q)s:1 0;:j~

><cOSs:1Q)

ttl>Q)1--1~ CJCl)

cOC\J

rl'rl (Y)1--1P. (Y/

c:r.:Q)1--1

?o'rlf:r......:::t 0 \0 C\J CO.,'. C\J C\J

JO a.rn '+B.ladura,L

Page 72: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

?. •rl;::1r-;,

s::'MS

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Page 73: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

UJ>.cd'd

Lr\

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H0~

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Page 74: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

Cl)

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Page 75: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

3.4 RLSULTS

Soil tempe r-ntur-e , a t 10cm depth ,"~1d screen, rn.;x , and min. a.i r

temperature did ~ot vary substantially between yc~rs (Figures 3.3.3(a,b)

.ind 3.3.2(_>, b) ). However, r~i:lf:tll (Fig.3.3.1) did differ be twe e n

ye ar s w i,th :J. no t i ceubl y dry period, in the men ths of J une v.nd July

duriLg 1979.

Dependi~~ u~on the results of 197), Lhere were only 2 treatments

in 1980 i.e. p~:)333 aprayed on 23rd H,,-y, 1980 and control, arid Cl. further

treated plot 1:1:;S p.lan ted at closer syCl.ngs (61X20cm). Gener-al, growth

of the crop was more rapid in 1980, in the absence of a dry period.

Stolon ,lYldstem growth

Before tuber ini t i at iou , .irnmedi .ite Ly .if t.cr s:)r1yin(; the pp333,

total dry we ight of the s to.Lo.is cmd to tu .rumbe r of stolons were in-

cr-eaaad in tr-eate d plots. ;~\'t the Lmpor t.rn t result 'd-,S th,.t, whee

plant growth reguL~tor \,/",_sspr.lyed be f'or-e tuber .in.it i a t ion , the per-

ce ntage of stolons out of tab} dry weigh t 1;J:lS higher in tre.rted plots,

for exampLe ,~dter lt4 d.iys of p.lan ting, in c.iso of G5 pe r centuge of

stolons out of to tul, dry we igh t 1.~j_S 3.55 compar-ed to 1.85 for control.

~imiLJ.rly il: 1980, "48 d:.ys .if te r pLu,ting the pe r cen tage of stolons out

of total. dry we ight w.rs 4.35, while .in con trol it W:JS 2. GO.

~o effect of np333 wus found on the cumber of stems, nor the

"'x"l b ' (~~)", t i ; ary r-ancues hD, present. Howe ver , stem exteus ion '.v;).5 decreased

in both yeu-s ir, t.re.rte d pL1: t s , Figure 3.Lr.1 ahows tbat pp333 decreased

the aver-age Length of t.he m.iinatema iJ_TId t.re r-espor.se V/,:J.S substantially

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affected stem he igh t (19?9), rr ther- where s~ri~yi?1g Vi~lS done earlier in

the season for example, G5, the rite of stem extension, later in the

ee aacr; was not much Lower th.m control. This m.iy be e xp.lai.r.ed by

lower up take of the cnemi c.cI due to dry ".Nec( there The p.i ttern of

?p333 effects on .rve rrgc .inte r-node Leng th 'tI,_,S s imi.Lar to tha t for stem

Le ng th (Fig.3.J+.2), indic .t ing Uut"l1 r.n ter-nodes i~rO'.-vingaf te r- <lp[.)l-

Lcu t i on were shortened. 11, 1979, Le.ive s for the Lus t two growth

'-uDlyses were not ccun ted, thus .ive ruge if: te rr.ode Leng th could not be

ce.l cul.a ted , Dat a for tote I dry weigh t oflbove ground stems (AGS) is

presented in Figure 3.4.3 und wus reduced in both years by the growth

reguLato r , Growth of under ground stem (UGS) WiJ..S not '_tffected, re-

sul t ing in d.i f'f'e r-ent io.l v.rl.ues for fiGS to DGS rrt io , on weight basis

(Fig.3.4.4 ) :;,Ed ',;ns higher in control ir: both year s ,

3.4.2

Decrease in stem Le r.gth v-u.s due to de cr-ec.se ir. internode length,

thus led cumber we re uo t .rff'e c ted , During both ye~<rs the foliage was

a darker shade of green following uptake of pp333. The area of individual

Le.ave s ',vas de cr-eaae d by pp333 ~lr.d the effect was more pronounced in

1980.

The growth of-leaves in control plots wus lower in 1979 comp3red

to 1980 (Fig.3.4.5 ), and , in .add i t i.on , up t.ake of pp333 may have been

lower under the drier conditions.

Irrespective of time of .ippLi.c.s t iou , p 1979, the DD333 - treated

plants had 11 Lower Le.if .ir-e., .inde x since le::fclre:1. aLUlt-1 was decreased

and pIan t spuc ing 'vias the same :,l5 the control. In t r-eo tmen t , G5, plants

h:,d >. highe r L!,I th .n , G18, 1 Iter ill the se.rson did ol onts s trr ted

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growing out of' its effect. Since 1e::f':"'8',. oer ;"ll:~nt '..!:'s expected to

be decr eased , the tr-eate d :Ire S in 1980 wer-e C11nnted t closer spn.cing

in order to ut iLase vrlI the .rv.ri Lobl e «er i .1 sp.ice , for compari son of-1

productivity 'ireJ unde r full c.inopy cover. Therefore,lthough the

:Jver'~ge Le.rf .ir-e.. ',Jas decr-eussed by :)()333, the Le.rf .u-e.. .i r.dex 'N'lS

s imi.Lar to that of the control dur i.i.g the 1980 experimen t , except ,'jt

the peak , wher; con t.r-ol LAl wen t uo to 5 .urd np333-treuted ,fBB 3.5.

Ir: 1980, iJp333- tr-eated plants senescedJ few days e .u-Li er thm the

control, resulting in ~ more rapid decline in LAI.

Changes in Leaf "rei). reflect ch.inge s in Leaf exoane i on Ln two

dimensions but ac tua.l leaf Growth occurs ill three dimenei ons , 'rhere-

fore ,'il though ')1)333 decre::sed Leaf »r-e.. the differences in Leaf dry

weight were not in proportion, resulting in differenti:J.I values for

specific Le.if ar-e.:, Specific Leaf .-lrel \oJdij decr-eise d , in both years,

by pp333 and the decr e.xse \r1::lS evident throughout the growing season

(Fig.3.4.6). It (pre ired thltPD333 h.id .increaaed either leaf thick-

ness, ,cellular density or stored carbohydrate, or a combination of these

factors. In the case of G5 (1979), the effect on specific leaf area

seems to diminish later in the season.

, 3.4.3 LiGht interception

Figure 3.4.7 shows that the nroportion of ?AR intercepted by the

crop canopy WD.S altered oy np333 tr-ea trnent , In 1980, light interception

vns s irni.Lar for con trol and [In.333 tre -J. tments be tween ?O - 105 d3YS after

planting (middle of the growing seIson), during this period LAI was,..

higher than 3.0 C_'.ndlight .inter-cep t i.on "'FiS independent of LAI at indices

of more than 3.0. LlU hard Ly r-e.rched 3.0 .in 19?9"-nd consequently

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t re a tment differences wer e eviden t thro1ii;hou t the growing season ruther

than a t no.rticul:ir times.

Tuber growth :~d development

The effect on tuoe r Dumber dur i ng 19?9 '-"';S v.rr iobl e (Fig.3.4.8).

G5 decreo.sed tuber number, l;-;~'y be due to very 101': Leaf ar e. index .rt

the time of tuber ini t i:. t io.i, AI though G1 S sligh Uy reduced LAI, but

Lncr ease in ava i.LabiLi,ty o f :-;_ssimil:ltes by de cr-eas.i.ng the stem gr-owth ,

increased tuber numbers. In 1930, tuber number in pp333-treated nlot

were affected in two wuys: firstly, due to closer srucing, there was

higher Le af ar-e« index :It the time of tuber .i.ni t i at ion , .md secondly,

due to r-e t ar-dut i on of stem;rowth more .lssimilutes were :wailable for

tubers. 'I'hus tuber numbers were i.ncr-e.cse d severalfold (Fig.3.lt.8).

Since totill light interception w~:..sno t much different,'lJ.though stem

growth w~s reduced, but there were not enough assimil~ltes available for

i)_11 those tubers to gr-ow , thus differences a t f i.n.r]. h.ir-ve s t wer-e re-

duced.

In 1979, 818 Lncr-eaee d tuber weigh t , es r-Ly Lr: the season by making

more aas i.miLate s uvcliL.lb=-e. Since Leof '"re:' index was reduced by a.LL

the p-;>333app.l i c.i t ions (1979), thus f i.nc.I tuber yield \'IUS higher in

control (Fig.3.4.9). In 1980, pp333-treuted plot was plrulted at closer

S~')acing, thus total ligh t illterce9ted by the canopy during the growing

seQson was the same, If then total aas imiLute s ava i.Labl e wer-e equal in

both tre ctmcn t s , p:::J333by r educ ir.g stem gl'o\'1th and m.iki.ng more assimi-

lates available for tuber .Gro·",th, r-esu l ted in 16ib higher tuber yield

area -1 over control (Fig. 3. 4.9) itt i'il,:'ll h.rrves t ing•

. Although taL,] tuber y i.eLd is .i.mpor t.mt , the distribution of tuber

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, .'i _)

sizes making un the to t.xl yield muy :~}so be of importence in practice.

In both ye ar s , the IY:_J333-tre·,ted crop h·'d.l. highar- pr-opor t ion of tubers

in the 35 - 60r.;mr' ..G1ge th"L the control (?i;::;.3.'t.10.:', b).

3.4.5 rrot:;,l dry r:kltter :)ccur.1uLttio!:.

,Roots were not collected ·.llld those pr-ese nt OL stems or stolons

were removed and di.ac.cr-de d , thus t.ot.i.l dry weight (TU:I) presented is

exc l udi.ng roots. Lat.e r in the ae.ison (er:d of July) Le cve s started to

f a l.L off ;,nd were not collected from the ground. The TD\'!uchieved in

1979 was sUbstanti;lly lower ellD in 1980, reflecting the l~ck of rapid

srowth dur ir.g the dry period in June and July of 1979 (Fit~.3.4.11).

In 19'79, P?333 :'.t:Jll t i nes of l:)plic·tion reduced TJi"i, since p.lan ts

were shorter .r.d took U~)lessler:i.:J sp.ice , In 1900, np333 treatment

-1':J':S grown .it closer SraCi;lg to de te rmi.ne if dry m.it.ter production are a

could be .incr-er.se d, 'I'here 'tJ,";'8 very little d.if'f'er-ence i~l dry weight m-2

wi th l)P333 .md closer ap.ic ing cor.par-ed to trie cor. trol (Fig.3.4.11).

1',1though the gr owth regula tor 1110wed closer ap.rc ing IIi thout visible

crowding effects, inter-pl~nt competition for w~ter, nutrients and

soil space Iil:1Y have influenced the crop pcr I'or-m.u.ce,

~Given thut t.otal, dry mat te r pr-oductior: 'lre:;;-1 Ions slightly de-

creased (1979) or airniLar- '.'lith closer SP'~Cil:S (1,)80) it was of obvious

imporb.nce to determine the distribution of th.rt dry m.it t.er- since tuber

Yields i~re the commercial product. Figure 3.ll-.12 Lnd i oa te s tha t in both

ye~rs there w~s :J. higher proT'OrtioE of tot 11 dry wei3ht :J.lloc:J.ted to the

tubers ire the pp333 tredment. The r'edi s t r i.cu t ion of dry weigh t ents

more evident e~rlier in the se~soll, probajly bec3use tuber initiution

occurred 4 - 5 d.iys c.u-Li er w i th pp333.

Page 80: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

110

88

66

22

66

22

Key: D,G5; O,G18; 6. ,G2; v,G16; o,Control

1979

o~ ~_June July August

Key: D,PP333; o,control

1980._ B'

$if-

/////

¢//

/ __-----Brn --~-r: --a-----.~----

~-.

O.....__--.,.----.- ..-~- ...------__"T-------June JulyMay August

Figure 3.4.1 The effects of PP333 on average plant height.

Page 81: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

55

46

3'1

28

19

s 10o

....c:+'tID$:lQ)

rl

Q) 55'd0$:l!-iQ)

+'$:lH 46

3'1

28

19

1979Key: 0, G5; 0' G18; o,Control

..e--------E)./

././

./.f2f

-:-:

,/,/

G---ff

~--

-r--------· -·--·--------r·------15 15

June July

1980Key: 0 , PP333; 0, Control _--8--J2j--

./ -

-:./

,/,/

./~/////

¢/

_-_-B--.-----_£]

10....__-----,-----.---·---i----r--------.15 15

Figure 3.4.2

May June July

(main stems only).The effects of PP333 on average internode length

Page 82: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

i1

I135J

III

102~iI1

58J

Ii

341oL:-------- ..-----r...------..---..-------..-..-..-.--.-_._--,._--_._-----

170...,I 1919!

June

Key:170_,

!I,I

iI

135~

1980

I

102i1I

III

58J

fd- __

/ p' ------=:-=-.:E1//

//

~-/

/ ~c:--_A__---v

/

July August

.Q/' "-o ,PP333; o,Control /' /' "-

/' "-cp ""-/ "-/ ""-/ ~

/////¢///

Mayr .--.._._- -.-. 1- _ --- .'_--'_"

June July August

Figure 3.4.3 The effects of PP333 on dry weight of aboveground stems (AGS).

Page 83: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

O,G5; <>,G18; A,G2; V,G16; °,Control

055

042

030

018

././cp.

/

B- -_../ -

././

005(/)

June July Augus~0:::><,(/)0<:t:

0, PP333; o,ControlX101 Key:06'1 1980 Jd-_

/ -- -- ~B/

055 ///

¢

042 ///

q5030 I -~

I --~I

018 I

#

005May June July August

Figure 3.4.4 The effects of PP333 on the ratio of total aboveground stem to total under ground stem (dry weight basis)(AGS/UGS).

Page 84: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

X'10 1052-

1Ij

o ~2)

031JIr

010

Key: O,G5; O,GlC); .6.,G2; V,G16; Oj Cont.z-oL1979

_--0,0- --

/ 0---------__-4//

Io 00.J----------r--------.--.---- 1

_. _

~/ \/ \/ \/ \\

V\ \o

/¢/

)/J-:

O 0'o 0 ----,---------.- -------- .---r----------.--.----r---.----------

May June July August

H

j

X101

o 5.2joi2

I

010

June July August

Key: 0, PP333 ; 0, Control1980

Figure 3.4.5 The effects of PP333 on leaf area index (LAI).

Page 85: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

1050June J.lly August

rlIt10

C\J

~ 30 SOlKey: O,PP333; 0, Control

1980 _G.

~ / <,<,

Cl) / <,

3010J / <, _-0-/ <, ---e--/

I ¢

20 '01 // 7----

20301

/ /)OJ// \

/ ~

1090

Key: O,G5; O,G18; ~,G2; 'V,G16; Oj CorrtroL

1979

10 50-'-------r---------------l---------- ---------r---------May June July August

Figure 3.4.6 The effects of PP333 on specific leaf area (SPA).

Page 86: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

20June July August September

s:::0''';+> 0, PP333; 0, Controlp, Key:~OO1-< 1980QJ+>s::: /',.;

~n / ~~ 8{' » \p., qf e-- \'* I ~

I <,

68 I"(\)

\I \ \I \52 I \

I c.'9.<,

¢ B

36 //

0

20June July -r- August Sept.

Figure 3.4.7 The effects of PP333 on photosynthetically activeradiation (PAR) interception.

100

8{'

6R

52

36

Key: 0,G5; 0,G18; 6.,G2; V,G16; O,Control

1979

Page 87: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

225

180

135

90

45

C\JIEl

~ 225OJ

~s::~OJ 180~

135

Key: O,G5; O,G18; ~,G2; v,G16; O,Control

1979

o -I--------------------r------June July August

Key: 0, PP333; O,Control1980

Figure 3.4.8 The effects of PP333 on total tuber number.

Page 88: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

till

<) , G18; 6. , G2; V, G16; 0, Control s::Key: 0, G5; .,-i

+>-1515 Ul

.-l (l)al :>-

1979s:: I-i.,-i alrx..s::

I1()C () !(~u, -1

0i!

9L:5JAPI

II

530JIII

3-15_1

0 0-C\J --- ---.--------------I June AugustSeo"+>..ceo

.,-i(l) Key: 0, PP333; o ,Control~ 1575

l>. 0I-i 1980rc::J

I-i(l)

0.o;::J 12608 e

././

./9{'5 '/

y~

-~'/

~_/

:/10

././

./-:

0~_H_=a__=-=-----=---0-----1----------------1-----------June July August

630

315

Figure 3.4.9 The effects of PP333 on tuber dry weight.

Page 89: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

80 •

40

•60 •

••

20 ••o

G18

o 0

o 0

o 0

o 0

•••••••

G2

••••••• •G2

•••

o 0

o 0

o 0

• •

G16

•••••• •

Control

o 0 0

G16

••

o :::>

• •• •• •••

o

92 daysafter planting(DAP)

•••••

G18

•••••

o

tJ 0 0 0

o 0

Control

•114 nAP•

••

0>60mm

[!] 35-60mm

~<35mm

G>57mmo 32-57mm

Final harvestingFigure 3.4.10a The effects of PP333 on proportion of tubers In

size grades (dry weight basis( (1979).

• • •

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. ..oj.) 0... ..__"--_-'--~-I

~'r-!Q)~

t>'0

HQ)

.g 70oj.)

~60

Control PP33380 - •

•••••• I---

• • ...

60 -

• •40 -

• •• •20 -

• ••

107

Control PP333

••• •• •• •• • 1

••

r--- •• •• •• •

135Days after planting

Control PP333

0

0

II III 0 ~

40

0

0

C

0

0

rrrn- 0 ~

20

0,

Final harvesting

G >60nun

G 35..,.60nun

8 < 35nun

OJ]] >82nun

D 57-82nun

0 38-57nun

I~~~d < 38nun

Figure 3.4.10b The effects of PP333 on proportion of tubers ln sizegrades (dry weight basis) (1980).

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-1500Key: O,G5; O,G18; ~,G2; V, G16; O,Control

1979

1200~1i,I

q'JIJ!'-' -II

II

600

1300~

I

oL --r-------------------- ---- ---------r----- -------------July August

sa--////

June

C\J 1 c: r'oI ... ,~I J

S!:lO~

::;:~ 1;~OOE--l

Key: 0, PP333; 0, Control

1980 -:/"

././

./50

//

//

/¢/

,?

300//

125-:

/'J2fo _-0'..J.---'=-__ r---------------r-------

May June July

900

600

August

Figure 3.4.11 The effects of PP333 on total dry weight (TDW).

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80

'15

60

45

30

15

0

~E-i'i-i0

+J;::J0

10-< 80_w'§+J

*'12

54

18

Key: 0, G5; 0, G18; .6, G2; "'V, G16; 0, Control

1979

-'-----,------------------;----

June July August

Key: 0, PP333; 0, Control

1980

Figure 3.4.12 The effects of PP333 on percentagesof tubers

out of total dry weight (TDW).

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3.5

ion of the grm'Jth ret;ulator, ;:;:)333, I!ere:ou!ld ill two ye.ers of expe r i »

ments , Ilowe ver , the m,.lC}li tude of the r-e scor.se 'du.s lower in 19'79 probably

be et.use of drier cond i t i ons (FiC.3.3.1). '';:'11eLn ter-ac Lion with soil

moisture pr-o b.ib.ly r-eLi te s to upt,kcmll eubae quen t tI"111s;Jort in the

xylem r;;.ther t.h.m d i f'Fe r-er.ce ill grO'.:th recu.,l tor .ic ti vi ty ~ ~.

'I'o tal, dry m.rt te r pr oduc t i on \J:)S sliGhtly de cr-e.ase d in 1979 irresoect-

ive of the time of :,,~)'plicitior: of the grm·!th recul'-Itor and it may be

tha t h.id up t k e of prJ333 beer. gre,:ter tho n t llrcer effect would have

been f'ound , II: 19'79, there '.ns :i de cr-ease in dr-y weight ,Lmt -1, and

sir:.ce pl ant der.s i ty ':/:lS the s,'.me ·'.S for tile control, then cC decr-ease in

;->roduct ivi ty ui. i t ,~rcl-1 Hi th rn333 rcsul ted. Ilowevor, since individual

9limt size VI:lS de cr-eaae d the po te nt i iL exi a te d to irlcre:~se plant density

w i thou t sufferill,';'" de trimer: t,~l degree of'lLm t compet i t i.ori, The experi-

',"eigh t un i t-1

~re:l was not decre~sed at closer plant s~acing combined

w i th p)333.

'I'otal. dry mi tter produced by the c.mopy depends on the light inter-

ce_?ted by the C~,llO~)y. Si!1ce L,~I \'I',ISreduced by ~,ll the p_)333 treatments

L1 1979, Li.gh t intercented 'ler unit 'lre:, \'IitS lowered ,md this resulted

in lOi-Jer tot::,J dry mltter in thc tre·,ted~)lots. Ir: 1980, LAI was

Blightly reduced durinG the ~e(J: but L1 W~6 ~ot ~5 LA! in pp333 during

th:J.t geriod ~·!.~sover 3.0, for liCht :terccotio!1 ioDS indepe:ldant of LAl

:,t it:dices of more th::L 3.0. 'ire:;,tcci·ll.lL:S interce'lted more r:ldiation

e:::.rly in the se::501: wher; so;_~r rJ,di.:...tior: 'IJ ..fj hig;}, :md so· toL'.l dry

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An add i t i one.L f'ac to.r , 0 ther than solely the degree of pp333 uptake

which m~y hiJ.ve influenced the 1979 results is the possible interaction

between the phys i.o'log i cru s t a tus of the pl;"r.t and "·:~tter stress. The

control p.Lunts ere'cl more slowly ..md yielded less in 19'79 th m in 1980.

The crop was visibly .~ffected b;y dr-ough t Lr: 1979·,nd the calculated

net aas i.m.iLat ior. r-ate C:AE) de cr-e.ised dur ing the dry »e r i od , whereus no

de cr-ease in i·.APW:J.S found i:; 1980.0il333 h.id little effect on ::AR in

1980, or in 1979 with the exeeution of the dry period. During the

d::-iest period (second week of July), the control lJAR fell from 7.0 to

-2 -1 81.1g m day while the !JARof G5 and G1 treatments were maintained

-2 -1at 3.0 and 4.5gm day respectively. Therefore, the efficiency of

dry matter accumul.u t ion \·us chr-e e to four times h ighe r in the pp333

treatments during the drought period. Other growth regulators such as

eee have been reported to increase toler(lDce to \Vater stress (Teubner,

1961) and the effect may be r-el.a te d to changes in physiological status

iL response to Cl general r-et cr-dnt i.on of haul.m extension growth.

Reduction in notato haulm growth and some degree of redistrubution

of dry matter to the tubers has been reported f'o.lLow ing vrpp.l i.ca't ion of

ccc (Dyson, 1965; Gunase na .irid Harris, 1969, 1971), and 139 (Humphries

and Dyson, 1967). t,Hth the'1.pplication of pp333, haul.m gr-owth was de-

cr eaaed arid there vldS some redis_tribution of dry mat te r to the tubers •

.In 19.79, the pe r-cen tugea of aas imi.La tc s diverted to the tubers were

lower in G5 ., compared to G18. This was probably related to a greater

uptake of the chemicul by the G18 treutment us LAl \Jas 0.7 at the time

of its application while G5 was spr-ayed 4 - 5 days after emergence and

soil uptake must have been very 10\01 as weuther "1:':lS dry in that year.

Percentages of assimilates diverted to the tubers were higher in treated

plo ts and since sp::cing w::.s the aame for -;;reated :,lS \\Iell:1s controls

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which resulted in lower LAI in90333 tr-e.it ed plots JEd so the finu.l

tuber yi.eI d w:~s higher in the con t.r-o.l, In 1980, the per cen t.age s of

aas imi.Lates diverted to the tubers were much h.i.gher them in the control

and differences were evident throughout the season , :probc~bly related to

more uptake of the cheml.c al. in a wetter season. SiI,ce t.ota.L light

intercepted by the treo.ted?lots ,·,us the s;',r.Jeas in the controls, final

-1tuber yield wus 16 per cent higher urei:. th.in con tr-ol , The foliage

was darker green, as occurred with CCC (Gifford and EoC/rby, 196'l),

and the leaves were either thicker or denser. Chlorophyll content per

unit are3. was 25 - 35% higher ir.. leaves from pp333 treated "Olants in

an ad jacen t experiment in 1980 (Hcl.ar-en , pers. commun, },

The higher proportion of dry weigh t f'our.d in t he tubers of chemically

manipulated plants is useful only when present in the appropriate tuber

size. pp333 resulted Ln :_,higher proportion of tubers in the size

. range of 35 - 60mm. 'v/hen ni trogen w:,s app.l i.ed early Ln the se ason , LAI

was higher at tuber initiQtion compared to late application or no

nitrogen. Since LAI was high, the assimilates available for tubers

vlere more and this resul ted in :.l higher number of tubers (Gunaaenc and

Harris, 1968) • Pratt, et al., (1952) , have shown that irrigation during

the period of tuber set increa.sed the yield of the crop by increasing

the number of tubers set, whereuo? when w:lter was applied later in the

~euson, irriGation-had the effect of increasing theci.verage tuber size

rather than tuber numbers. Photosynthetic efficiency of the canopy is

higher when soil mosture content is high U~oorby" et :11., 1975; Legg et

c~l~, 1979) thus Lr-r i.gat i.on a t tuber set increased the ava iLabi.Li ty of

&ssimil~ltes for the deveLoping tubers, which resul ted Lr. higher number

of tubers. The mechun iam unde r Ly i.ng the pr eser.t exper iment may be re-

lat~d to dry mutter redistribution at an early stage of growth. For

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4'7

eX9J11ple, the number of tubers present ',/J.S s irniLar 'out it may be that

the amount of 'lsimil::l.tes diverted to those tubers w,',~sinitia.lly higher

in the pp333 t r-eatme nt , If et max imum r a tc of gr-ot ...th for individual

tubers is iJ.ssumed theri the add i t i onul .ias inri.Lat e may h.ive stimulated

growth of tubers whi ch o the rwi ae woul d not have GrovJrl at tD~l.t time.

Since more tubers woul.d the n h.we CA t tr ac ted more assimi1a te produced

subsequently, i.nd i,v.i.dua.I tubers wou.l d h.rve h.id f'ewe r- c.ssimiLltes due

to inter-tuber cornpe t i t i on, The resul t of such ., me charrism , r'e Late d

to assimilate ~v~il:Jbility ~t particular growth stages, would be to

produce more tubers within the medium size n.lnge,'~s was found with the

p:_)333 treu tmen t , Encr-e aae ill Lumber of tubers hits also been reported

when eee (Cunuaena and IlJ.rris, 1969; Gifford and Hoorlll~*,) 1967), or

ethrel (Gar-c ia - Torres and Gomez -Cumpe , 1972; Hurti et ::.;.1., 19?8;

Bil.nerj;ee et a1., 19'79; Pe r-uma.l,et :3.1., 1979) were spr-ayed a t tuber

initiation phuse.

Ifenk ....e'.:'.nd Allen \1978), showed that at Lowervp.Lan t density (24960

...., h -1) 11 t b d 1 d f th .. t i t d '0 t h i h~Uoers a a . u ers eveLope ram .ose 1nl la e ut et 19 er

-1 1plant density (74880 tubers ha ) about 1000000 tubers ha- were

initiated (June) but only 78000, developed (August). In 1980, all the

tubers Lni t i.e t.ed in pp333 tr-e.i ted plots did not develop. It may be

,explained 'th.rt ::l t higher p:Lmt density LAI .rt tuber initiation was very

high, thus more tubers were .ini t ia te d , but the r-ate of increase of LA!

Has 10\'; due to .inte r-pLant cornpet i t ion, Thus all the tubers which had

ini tiated could not develop because the aas imiLa te s ava i.Lub'Le following

tuber initii.J.tion \'Jere not enough for:.J.ll those tubers to develop.

In addition to any benefits from manipulation of the physiological

processes within the crop, the altered canopy structure may have agro-

nom:tc benefits under particular circumstaLces. For eXdIDple, the canopy

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48

micro-environment may influence the build-up and spr-ead of pathogens,

and the decr-ease d canopy height may decr euse the degree of lodging

and rotting of stem tissue which often occurs under wet conditions.

In 1980 lodging occurred in cor.t.r o.l plots .md rotting of stems W:J..S

observed, but rotting of stems \'hlS more in Experiment JTl~ where stem

-1number-a area were increased by using large seed. 1:0 lodging occurred

in the sprayed ([')9.333) l)lot.

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4. PHYSIOLOGICAL AGEl SPROUTING

TECHNIQUE AND SPACING

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4.1 LIT.E:RATURE REVI£~'I

4.1.1 Physiologic~l ~ge

It is well known thn.t for a period Immed i.ate.Lyafter harvest, no

appr-eciabl,e sprout Growth "/i:'..lb":e place on seed tubers, even when they

are stored in conditions ide~l for growth. The length of this period

of inactivity is referred to as the 'dormant' period (Burton, 1963) and

it ranges in normal storage conditions (100C) from 5 to 14 weeks after

harvest. The dormant period is largely determined by variety (Krijthe,

1962; Bu;rton, 1963; Bornman and Hamme s , 1977; Reust, 1978) though not

related to mo.turity classes i.e. early varieties do not necessarily

start growing before main crops (Emilson, 1949). Hany features of seed

crop husbandry also affect the length of the dormant period e.g. time

of planting (Jones; O'Brien (both quoted by Ali, 1979); Allen et al.,

1979); site of production (O'Brien and Allen, 1975; \'/urr,1978b); time

of haulm destruction of the seed crop (Hutchinson, 19780.; Vlurr, 1978b);

time of harvesting (O'Brien and Allen, 1975; Toosey, 1964 ); and state

of maturity of the tuber at the time of harvesting (Krijthe, 1962;

Hutchinson, 1978b; \vurr, 1978b). Temperature during storage also affects

the length of the dormant period (Schippers, 1956; Sadler, 1961; Headford,

1962; Burton, 1963; Short and Shotton, 1970; \Vurr nnd Allen, 1976;

Bornman and Hammes, 1977; Hutchinson, 1978b; Allen et ale, 1979; Jones

et al., 1981). \oJurr(1978b) stated that differences due to date of

defoliation of seed crops on sprout length were due to its effect on

dorm<lncy br-eak and 0'Brien and Allen (1981) regarded all seed stockswh i ch do not sprout i.e. until dormancy break as the same. Thus the

period of post dormancy break is important for this may affect the field

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50

growth.

Occe buds huve begun to grow however, the e~vironment in the

store, eape ci al Ly the tempero.ture, becomes extremely important in

determining subsequent sprout growth. The gr-owth of sprouts is pos i tively

related to temper a tur-e over the r-ange from u minimum of 4°C to 25°C

(Sadler, 1961; Headford, 1962; Morris, 1966). Growth rate of sprouts

at 300C was 10\'1 due to deatl: of the sprout~.I.pices, Lut.er dea th of

sprout apices occurred a t 25°C also (Head f oz-d, 1962). Horeover sprouts

produced 0. t such high temperatures are bulbous in shape [mel restricted

at the base where as those produced at lower temper-atures are more

firmly at tached (Davidson, 1958; Short and Shotton, 1970). Due to this

oreason temperatures higher than 15 Cure not usually used in the store.

A linear r-eLat iorish ip between total sprout gr-owth per tuber and temp-

erature accumulated over base temperature from dormancy break has been

reported by several workers, when tubers were stored in conditions

ideal for sprout growth immediately after dormancy break (;vurr, 1978b;

Ali, 1979; RaVJi, 1981). Toosey (1963) and l-lade c and ?erennec (1962)

described physiological age as the physiological state of the tuber at

any given time. Recently the research groups at University College ofV/ales and NVRS, Hellesbourne have suggested the measure of physiological

age as day degrees above a base temperature from dormancy break •

. (O'Brien and Allen, 1981; Ali, 1979; \vurr, 1978c).

Varieties differ in their rates of sprout growth Qt Q given temp-

erature (Headford, 1962; Headford and Ingersent, 1962; Short and Shotton,1970; Allen et al., 1979; Bod.Laender and Harinus, 1981) and thus may

emerge at different times in the field. Greuter differences in sproutlength at the time of planting have been found to affect the emergence

and tuber initiation ('rr), physiologically old seeel emer-ging and

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51

initiating tubers before the physiologically yOlli~gseed (Headford, 1962;

Toosey, 1963; Fischnech and Krug , 1963; Younger, 1975; Raquf, 1979;

Ali, 1979; Rawi , 1981). LAI .md total dry wei.ght a t the time of 'I'I

wer-e higher in nhysiologically young seed (R<~quf, 1'979; Rawi , 1981 j

Younger, 1975) though difference varied between years arid var-i.e t i.es ,

In the variety HomeGuar-d ,'l close relationship be-tween tuber yield and

physiological age has been found, tuber y ieLd being increased with in-

crease in phys io.Logi.cu'l :J.3e but the effect changed as the harvesting

VI'!S delayed and in some cases it became negative (O'Brien and Allen,

1978; Allen et al., 1979; Raquf, 1979). Decrease in tuber yield with

increase in physiologico..l age o..t later harvests was associated with a

decrease in LAI and total dry weight w i th increase in physiological age

(Raquf, 1979; Rawi , 1981), which may be overcome by decrease in spacing.

Ali, (1979) work ing , with the var i.e ty Desiree and Jones et al., (19811)

wi th the var Let ies Arran Comet and Desiree also reported increase in

yield with increase in physiological age but in these cases also the

effect disappeared as the har-ves t i.ng W:1.Sdelayed. !'·bjor concern in main-

cr-oo varieties is not the ear-Ly yield but the f'LnaL yield which may also

be affected by the total duration of the bulking period of the crop.

Younger (1975) found that physiologically old seed emerged earlier and

senesced earlier than the physiologicaUy young seed. But still if

early in the season LAI in the C:1.seof physiologically old seed is in-

creased by decreasing p.lan t spacing and thus making greater use of solar

radiation of th a t time of the year (Allen und Scott, 1980) then even if

it senesces earlier, the poter:tial exists for higher tuber yields.

Stem number is now considered us the unit of population in potato

(Allen and Bean, 1978). There are two types of stems which may emerge

ab()v.tground i.e. main and br-anch stems (defined in Appendix E).

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52

Differences in sprouting regimes cause differences i~ oroportions of

different types of stems produced by the seed tuber (Allen, 1978;

Younger, 1975; Bagley, 19,/1). So stem numbers ;..,s such may not give the

right Lde a of populat i.on but sizes of different types of stems have not

been studied.

4.1.2 In ter-cro')Ding

Ddf'f'eren t crops or var i e ties are mixed depending upon their habi t

of growth to ensure better light interce1jtion and extensive exploration

of the soil for r-emoval. of water and nutrients. Inter-cropping of

mai ze and bear.s is quite common in sever-c.I parts of tropical America.

(Pinchinat et ,')1., 1976). F'i n.luy (1974) reported that 98% of the cowpea

is inter-cropped in Africa. Ln te r=cr-opp i.ng of cotton .rnd summer onion

is practised in Egypt (Nasr, 1976). Over yielding of grains for mixtures

due to the early f'Lowe r i.ng arid m.rtur-at i on of one comnonen t than the other

has been reported by the Lntern.rt iona.l Rice Rese.ir-ch Institute (1974).

In potatoes inter-cropping may be useful when one component emerges

first and grows at the exoe nae of «no the r while the latter may have

advantages later in the eeasou, Schepers and Sibma (1976) obtained

higher yields in var ious experiments by mixing ear-Ly and late crop

varieties of potato. Smith (1978) obtained signific:mtly (P: 0.05)

higher yield by mixing Ar-r-an Comet and PentLmd Cr-own in ulternate rows,

compared to PentLmd Crown grown alone. Chowdhury (1980) also obtain-

ed higher yields, when Desiree arid Hujestic Here mixed v;i thin or be-

tween rows than the higher yielding monoculture. This increase in yield

Was associated w i th increase in Leaf urea duration. r'~ixing of different

varieties especially within the row may be only useful when the product

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53

is required for starch industry or for other uses for which a mixed

product is accep tab.l e , 'dhee s or-outed:'cld unaor-outed seed of the same

variety were mixe d al t.er-n.iteLy w i thin or be tween rows higher tuber

yields were obtained than the higner yielding rnonocu.;ture but the in-

cr e ase was higher .in hajestic th.m Oesiree and the effect al so varied

between years (Chowdhury, 1980).

4.1.3 Light interception

Potato crop growth has often been considered iL terms of the LA!,

leaf area duration and net ~ssimilation rate (Watson, 1952). A linear

r eLat ionsh i.p hus been reported between tuber yieldmd Leaf area duration

when leaf ar-e.r indices ubove three were·J.ssumed to be three (Bremner

arid RadLey , 1966; Bz-emner+and 'I'aha , 1966; Gunaseria and ILlrris, 1968 ;

Chowdhury, 1980), which implies that light interce~tion, or photosynthetic

efficiency, does not limit yield ut Lea f dTe'l indices gr-ea te r than

three. Published reports on light .inte r-cept i.on .in po t atoe s are few

(Scott and \-Jilcockson, 19'/8; Allen and Scott, 1980; Bean and Allen, 1981),

while crop growth rate has been related to radiation intercepted in

barley, wheat and augar-bee t (Biscoe:md Gallagher, 1977) and maize

(It/iUL.ullS et d., 1965).

'I'otal, gLobal radiation has "bee n four:d to be pos i ti vely correlated

'with the yields of several crops (Sibm,-l, 19?O)md potato tuber yields

have been related to total radiation during the growing season (Scholte

Ubbing, 1959). However, Leaf photosynthesis is easen ti al.Ly a wave-

length dependent with photosyn the t ic.i.l Ly active r-adiat ion (PAR) being

defined as radiation between 400 - (,OOmn(lkCree, 1972). Although the

ratio of PAR to lotal r;"dii..ttion appears to be reJatively insensitive to

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54

atmospheric fuctors, and when considering both direct :Anddiffuse

radi.at ion is often tuken :~sbei ng 0.5 (l'io;lteith,1969), it rn.iy be am-

portant to measure PAR int.er-oept ion withi n crop cur.oo ies, One f'~ctor

which may Lnf'Luence w i thin canopy meaaur-ements of total r id i.ati.on,

r-eLi tive to P,\R, is the transrniasi.onof w.rveLeng the ..bove ,/OOnm (Holmes

and Smith, 19'77; Scott etll., 1968), with the relative differences

being related to le~f canODY size und ch~racteristics. Puckridge wld

Ra tkowsky (1971) in whe« t .md Jeffersmd Shibles (1969) in soybeans

reported increases in photosynthetic efficiency of the crop canopy with

increase in LAI.Thus a study involving regular meaauremen t.s of sprout growth during

storage and various crop characteristics in the field along with regular

measurements of light interception may help in a better understanding

of the growth and development of the potato crop in relation to yield •

.',

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55

4.2 OllJ:i:CTIV':':S

Greaterphysiologic~ll age resul ts in .m .i.ricr-ec.aein the percentage

of total dry matter found in tubers but reduces LAI which ultimately

results in lower final yields. If LAl is increased by m~nipulating

the density then potential existed for h igher tuber yield ,it f ina.l

harvesting. ;."ith this in mind the experirnen t iD 19'79WiiS carried out.

Depending upon the resul ts 0btai.ned in 19'/9,wher-e p).1ysiologicalage did

not affect the emergence or senescence or tuber yield, but bigger seed

emerged first and ser-esced first, in 1980 it was decided to repeat the

treatments of 1979 (Exps, 1<'2,F3) and since different seed sizes emerged

and senesced at different times, in theory if they are mixed, then

potential exist for Lncr-ease in tot.a'l durat i.onof the crop, So Experi-

ment F4 was undertaken to investigate this nnd seed with greater differ-

ences in physiological age were mixed.

Emergence in 1979 may have been af'f'ected by the growth rate of

sprouts rather than physiological age or length of the sprout , There-

fore sprouting techniques were changed before plar.ting by keeping them

in the dark, to see its effect on emergence nnd further growth and develop-ment.

In 1980 cold treatment of seed took more time to initiate tubers

than apical or multi sprouted when counted from the date of 50'/0 emergence.

This may have been c.tffectedby longer days, as cold tre:tled seed emerged

later and day length increases in spring i:l.ndshort days do stimulate

tuberization (Chapter 2). Thus in 1981 it was decided to see the effect

of time of planting on early tuber yield aridother treatmen ts were in-cluded to have results for two years. Since emergence w.as not affected

by differences in physiological age of the seed (except cold) in 1979

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and 1980. It was decided to investigate minimum number of day degrees

above which emergence would not be elilianced(Experiment F6).

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r::' ):J(

l1ATERIALS AIm H£THODS

The experiments were carried out iJ.t the Uni versi ty of No t tLngham

farm, at Sutton Bonington, over the three years: 1979; 1980; 1981.

In 1979 (Exp , F1), soil VlUS sandy clay loam (Field 32) and in 1980

(Exps. F2, F3 an.dF4) and 1981 (Exps. F5'J.nd F6), it VliJ.S sandy Lo am

(Field 10 and 6 respectively). On receipt, the seed (details for seed

source in Ap-pendices: Bj Cj D) in general was handled in the same w~

as described earlier (3.3), except that in 19S1 seed was affected with

Rhizoctonia and thus Vias treated with Polyram (d:hthiocarbamate)

(100g PoLyr am dissolved Ln 50 litres of w,'J.terand seed dipped for 2

minutes).

4.3.1 . Experiment F1

Experimental design and or-acti.cal,details

There were 2 vari.eties, Pentland Crown and Record. Each variety

\Vas given two sprouting treatments (physiologiccll age), apical

(stored at 120C for 110 days, starting from 20 December, followed by

22 days at SoC) and multi (stored at 4°C for 97 days, starting from

.20 December, followed by 13 days-at 120C ~nd 22 days at SoC), and then

planted at 2 spacings between plants, 30 and 40cm. Thus there were 8

treatments, all combinations of 2 physiological ages, 2 spacings and

2 varieties. 27th December was taken as the date when dormancy of the

seed was broken ""stotal sprout length per tuber on the tubers stored

at 120C was about 3mm ('Ivurr,1978b). Thus apical and multi had 912.',

° .and 192 day degrees above 4 C from break of dormancy respectively.

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The seed sizes used in the experiment u.regiven in Table 4.3.1.1.

Table 4.3.1.1. The weight of seed sizes used (g).

Replicates I II III

Variety

Pen tland Crown 34.1 ~ 0.39* 61.6 ~ 0.52 +105.3 _ 1.31

Record +39.6 _ 0.59 51.5 :.0.'72 +63.0 _ 0.48

* SE, ce.Lcu'lated by weighing 40 tubers .ind.iv.idua.lLy in every case.

The experiment was planted on 1st. Hay 19'79in r:mdomized block

design, consisting of 3 blocks. Spacing between rows was 76cm. There

were 6 or 8 rows (depending on spacing treatment) of ~trnetres each to

accommodate 96 tubers per plot. -1Fertilizer at the rate of 1318Kg ha

(15: 15: 15; N: P205: K20) vias given before p.Lan t i ng, Other details

sueh a~: stem emergence; light interception; plant protection measures;

weed control ; rainfall and temperature etc. were the same 3.S those

described in Chapter 3.3.1.

4.3.1.2. Growth analysis

Nine growth cmalyses were carried out for the vilrietyRecord and

eleven for the variety Pentland Crown. Inaddi tion Cl. few plants were

harvested from specific replicdtion (as size of the tubers used affected

slightly emergence) to have Cl better idea of tuber initiation. Hethods

of harvesting and laboratory pro.~edure, were the same as those described

in Chapter 3.3.3 (2 plants were harvested at each growth analysis),

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except that foliage was not cut off the ground in the field rather it

was harvested along with underground parts 3.lld separated into main and

branch stems in the Labor-at or-y, to study the contribution made by

different types of stems, par t i.cuf ur-Ly the LAI. 1:1 general s tat i s t i cal,

analyses were done as f'nctcr-LaI randomized block design, thus residual

degrees of freedom (RDF) was 1L~. But for studying different type of

stems analysis was done :,s a split plot design. RDF for split plot

analyses is given on the Figures itself.

Sprout growth during storage

Length of the sprouts on 40 tubers (10 per truy) each of the 3

sizes (Table 4.3.1.1) and of the two treatments (apical and multi) in

the both vcr-i.et ies was measured on: 23rd J..:tn.; 7th Fe bs ; 13th March;

11th and 28th April, in the case of apical and 11th and 28th April in

the case of multi.

4.3.1.4 Measurement of soil water content

Volumetric soil wate r content was determined at weekly intervals

be tween Hay and October for al.L the 24 plots using a modified version

of the l..Jallingord _neutron probe (Bell, 1969). Essentially this consists

of the emission of fast neutron from a sealed radioactive source

(some Am/Be mixture) and a count of the density of the cloud of slow

neutrons resulting from collisions with the hydrogen nuclei in the soil

wnter. Aluminium acceas tubes were installed ill the furrow about one

,', metre inside from guar-d pLant , Soil profile was monitored to a depth

of 100cm at 10cm depth Lnter cd.s, 'I'he dute of ini tic ...l extraction of

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Go

water by roots for individual soil horizons was determined as described

by NCGowan (1973).

This was as described eGlrlier (Chapter 3.3.3)

Exnerimer,t F2

Experimental design and nructical details

There"'fJ.ere9 tr-eatmen t.s , details are shown in Table 4.3.2.1. The

variety used was Pentland Crown.

Table 4.3.2.1. Details of experimental treatments.

Treatments

Apical

Hulti

Cold

Ml1 25, A+l'l36,MC 25, A+C 36,N+C 25, M+C 36,

Details

Stored at 120C from 28.11.79 to 12.4.80(both days inclusive)

Stored at 4°C from 28.11.79 to 26.3.80and at 120C from 27.3.80 to 12.4.80.

Stored at 4°c from 28.11.79 to 12.4.80

+ = two sprouting treatments mixed byplan ting a.Iternn tely along the row. 25and 36 is the spacing in cm w i thin rowfor that particular treatment.

N.3. Apical, multi and cold were planted ot 36cm spac.irigwi thin row.

All seed was moved to 8°c 24 hours before planting on 14.4.1980.

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61

14th December W:3.S taken as the date when dormancy of the seed

was broken (Chapter 4.3.1.1). Thus apical, multi and cold had, 972,

140 and 4 day degrees above 4°c from bre.ak of dormancy , respectively.

Seed sizes used were 116 : 2g in replicdion 1 nnd 62 :.0.98 in

replication>2 and 3.

The experiment was p.Lanted on 14th April 1980 in randomized block

design, consisting of 3 blocks. Spacing between rO\'JSwas 76cm. There

were 10 or 7 rows (depending an spacing w i thLn row) of 5.40 metres each

to accommodate 150 tubers in the case of 36cm nnd 147 in the case of

25cm spacing w i thin the row. Fertilizer at the rate of 1318Kg ha-1

(15: 15: 15; 1':: P205: Z:20) was given before p.Lant i.ng, Other details

such as: stem emergence; light interception; plant protection measures;

weed control; rainfall and ter.1peratureetc. were the same as those

described in Chapter 3.3.1.

4.3.2.2 Growth analysis

8 growth anal.ysea were car-r-Ledout and 4 plants were harvested

per plot at each growth analysis. In addition a few plants were

harvested from specific replicCltes (as size of the tubers used affected

slightly emergence) to aid the .iaseasmerrt of tuber initiation. Hethods

of harvesting and laboratory procedure were the same as those described

for Experiment F1 (Chapter 4.3.1.2). In generul statistical analyses

were done as randomized block design, thus residue:.ldegrees of freedom

(RDF) was 16. But for studying different type of stems'malysis was

done as a split plot design taking stem type within the plot as a sub

plot. Similarly for studying growth of 2 types of plants within plot

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62

was also done o.sspli t plot design , taki.ng the 2 types of plants in a

plot ilS sub plots. RDF for suli t plot ,mi_l_lysesis given on the Figures

itself.

Surout e;rowth duriLg stor-age

To study the growth of individuc.l sprouts, eyes were numbered, the

first eye (eye No.1) on the aui caL end .ind then working down towar-da

heel end systematicclly wi th last eye numbe r given to the eye closest

to the heel end. 40 tubers (10 per tray) each of the 2 seed sizes

(62g and 116g) illcase of apical and 30 tubers of seed size 62g and 20

of the 116g in cuse of multi were used for sprout measurements.

Apical sprouts were measured on 14 and 21 December; 8 [illd21 January;

4 ond 18 February; 3,17 and 31 Harchj 11 April and in multi on 1 and 11

April.

4.3.2.4 Heasurement of soil w:.l.tercontent

Soil water content W:J.S measur-ed for 18 plots (Replications 1 and 2)

as described earlier (4.3.1.4).

Final harvesting

This was as described earlier (Chapter 3.3.3).

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4.3.3, Experiment F3

Experimental design and Dr~"ctic',llde t.ri.Ls

There wer-e4 tre')'tments, combi.net i.onaof two physiological ages

(apical and multi) and 2 aprou ti.ng treatmen ts (L,st:~'1d slow). Det ai.Ls

are given in Table 4.3.3.1. Again the vari.ety used v/aB Pentland Crown.

Table 4~3.3.1. Details of expe r-Lmen tal. treatments

Treatments De t ai.Ls

Apical slow Stored at 120C from 6.12.79 to 12.4.Bo

Hul ti slow Stored i:.lt4°c from 6.12.79 to 4.3.80 andat 120C from 5.3.80 to 12.4.Bo.

ApicJ.l fast Same as anical slow, but covered withblack pol~thene sheet from 6.4.80 untilplanting.

}1ulti fast Same as multi slow but covered with blackpolythene sheet from 6.4.80 until planting.

N.B. All seed was moved to BOc 24 hours before planting on14.4.Bo.

15th December was taken as the date when dormancy of the seed was

broken (Chapter 4.3.1.1), thus apical cUldmulti had 964 and 316 dayodegrees above 4 C from break of dormancy respectively. Seed size used

was 77.5 : 1.26g.

The experimen twas plan ted in randomized block design, consisting

." of 3 blocks. Spacing between rows was 76cm and between plants 36cm •

There were 6 rows of 5.40 metres each accommodating 90 tubers per plot.

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All other details were the same as those described for Experiment F2.

Growth analysis

Six growth ana.Ly.sea were carried out and 4 ul ants were harvested

per plot at each growth ana.lyae s except for one (6th Jur:e), when 2

plants per plot were har ves t.ed, j\'iethod of harvesting and laboratory

procedures were the sume :lS those described for Experiment F1

(Chapter 4.3.1.2). In gelleri..tl statistical analyses were done as a

factorial randomized block design, so residual degrees of freedom

(RDF) was 6. But for studying different type of stems (mo.i,nand branch),

it was done as split plot design taking stem type within plot as sub

plots and RDF WJ.S 8.

Sprout growth during storage

91 tubers out of 12 t r-ays in case of apical, (46 slav; + 45 fast)

and 34 tubers (out of 4 trCl.Ys) Ln case of multi (14 al ow + 20 fast) all

wi th numbered elyes were used for sprou t me.rsuremen ts. In the case of

apical slow sprouts were measured on: 15th ancl.24th Dec.; 9th and 22nd

Jan.; 4th and 18th Feb, ; 3rd, 18th and 31st Harch; 10th April and for

apical fast, the first 9 dates were the same and after that these were

'measured on 9th, 11th and 13th Apri.L, In case of multi slow sprouts

were measured on: 12th and 18th Har'ch ; 1st and 9th April and for multi

fast in addition to these 4 dCl.tes sprouts were also measured on 11th

and 13th April.

In addition some tubers (6 per treatment before the st.ar-t of fast

and 3 after thut) were used to study the SDrout weight, 3 times before

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planting and 2 times after planting but before emergence.

Fin~l hnrvesting

'I'hi.s vias as described earlier (Chapter 3.3.3).

Exuerimen t FLt

4.3.4.1 Experimentc.lidesign and practical details

There we re 4 treatments: 336j S25j I3+S25j I3+S36, where:

B = Seed stored ut 4°c from 6.12.79 to 4.3.80 and at ~.2~C

from 5.3.80 to 12.4.80 'lndseed size was 204 :.5.7g

(316 d d b 4°C).~1.Y egrees a ove

C = Seed stored at 4°c from 6.12.79to 12.4.80 and seed

size was 62 ~ 0.9 (4 day degrees above 4°c).

+ = Two seed sizes (B,S) mixed by planting alternately

along the row.

25 or 36 = The spacing in cm used for that particular treatment

within row.

All seed waamoved to 8°c 24 hours before planting on 14.4.80.

The variety used was Pentland Crown. The experiment was planted in a

randomized block design, consisting of 3 blocks. Spacing between rows

Was 76cm. There were 6 or 4 roVis (depending on spucing within row) of

5.40 metres each to accommodate 90 tubers in case of 36cm and 84 tubers

in case of 25cm spacing wi thin row. All other detai.Lswere the same

as those described for Experiment F2.!II

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4.3.4.2 Grmvth ,:~Lll'';sis

Six gr-owth .mal.ysee wer-e carried ou t:md 4 ';)l:mts were harve s ted

per plot out OE 1st July ocLy one r-epl i.cu t ion w;_~sh.ir-ves ted and on 15th

July 2 replications were harvested, while on remaining dates all the

three r-ep.li.ca t i oria were hL~rvested. In add i t ion i.l. fe\-! plants were

harvested from specific replication to help with the determination of

tuber initi.J.tion. Methods of harvesting ~ld laboratory procedures were

the same '.l.S those described for Exper i.ment F1 (Chap tez- 4.3. ,1'.2) except

tha t differen t type of stems and ax iLl ar-y branohe s were no t studied.

St3.tistical an.d yse s were done :lS r andom.i ze d block design, thus resid-

ual, degrees of freedom (IWF) W:.J.S 6 where illl the three replications

wer-e harve atcd and 2 where, only 2 r-eo.li cn t iona were harvested.

Sprout crmvth during stordge

j~yes wer-e numbered ~,s described earlier. 20 tubers from 2 trays

were used in the cuse of B36 for spr-ou t measuremen te on: 12th and 18th

!--i,3.rchj4th and 11th A~)ril.

Finul harvesting

'I'hi,s \-:::lS the s ime '-cs described ear Li e r- (Chupte r 3.3.3).

',.

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6'7

Ex·oeri.:nent 1"5

EX1Jerimenbl design lind pr:~ctlcal details

There wer e 6 t re.rtmen t s i cold 6; :.,)ic.~l 13; /~+C 13; cold 16;

ap i cal. 22; A+C 22, whe re :

cold = Seed stor cd v.t L~oC frOIJ 1ltt:i Dec. 1980 until 24

hours before pl;m tinge

api cul, = Seed stored at 120C from 14th Dec. 1980 until 24

hours before 1JLmting.

= Ap'icol UY.d cold mixed by p.lcrrt i.ng ill ternately

1 tl (F.~ 1 -..; 1)a. eng 'le row ltJ. t.,/. •

6, 13, 16 and 22 are the d.rt es of pl.an t ing in the month of

April for -:nrticuLlT t.r-eatmen t s , All seed was moved to 8°c

24 hours before }l~~ting.

The var ie ty used ':IL.s DentLund Crown and seed size \-J:J.S52.8 -: 1.1g.

22nd December 'Ins t alce n as the d',tte when dorm.mcy of the seed was

broken (Chap t.er- L;-.3.1.1). SO .ipi.caL 13 .:t:ld:J.!_)ic:J.l22 h..id 892 and 964

d~'.y degr-ees above 4°c from dorrn.uicy ore.:,';( respectively and cold had 4

day degrees only.

The experiment was p.Lanre d in u r.:J.ndomized block design consisting

of 3 blocks. Spac i.ng between r ows wa.s ?6cm and be twee n plants 36cm.

There were 5 r-ows of 7.92 metres e ach :iccommod'.ltine; 110 tubers per plot.

Fertiliser the rate of 11251(g 11.]' -1 (1?:1'7:17; I'. P205: KP) givencl t .. was

before p.lan t i.ng , All other de ta i.Ls were the aame .:J.Sthose described

for Experiment F1.

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68

GrO\.,rth ,-uul vsis

Seven Growth :tu1.1yses wer-e c.u-r i ed out .ind for e xch , l~ p.Lants wer-e

har-ve s ted 1jer ol ot , ;'lethods of h.u-ve s t i ng .md Lrbo r.it or-y pr'cce dur-ee

were the aame )8 those used for Exne r i.ment Fll-. ,st:,<tistic'.'.: :1nn.::' yses

\'1').s done ::tS f'ac tor i a.L (2 d.i tc s x 3 trertmen t.s i.e. d£:lictl, rnu.l t i .md

1i.+C) r:mdornized block de s ign , Thus r-es.idu.i; decrees of freedom I"las 10.

S:)rout ?jrm'lth durin0 storlce

I';yes were numbered vrs described 'olrlier (Ch.ip te r '+.3.2.4) 29 tubers

out of 3 trays were used for sprout measurements on: 22nd Jec.; 5th and

20th JUl".; 4th~,;ld 1,?th Fe o., : 3rd .u.d 1'/Ll IL,rchj 1hth April 1981.

Exnerimer.t F6

4.3.6.1 Exnerimental design dnd prdctic~~ details

There were 10 treatments: 4D; 24D; 48D; 80D; 1230; 134D; 2320;

280D; 352D; 920D, wher-e numbers before 0 s tand for the number of day

degrees gi veY);bove 4°c 'from br e.rk of dorm.incy , Seed used was the same

as that used in Experiment F5. 'l'r-ea trnent 9200 \-Jas the aarne as upi.ca L

in E~periment F5. For the rem~ining treatments seed was stored at 4°C

until moved to 120C on: 3rd, 12th, 18th, 24th, 31st Mdreh ~~d 6th, 10th

Ll.nd13th Apr i.L, to ob t.ci.n the r-equ i red number of day de gr-ees mentioned

above. All seed was moved to 80C,24 hours before 9lacting on 16th

April 1981.

The exce r imen t \viJ.S pl.an ted ill ,\ r'.mdcm.iz.ed bl ock design consisting

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1980 (Experiment s: F2; F4)

0 • 0 •• 0 • 0

0 • 0 •• 0 • 0

0 • 0 •• 0 • 0

0 • 0 •• 0 • 0

1981 (Experiment F5)

• 0 • 0

• 0 • 0

0 • 0 •0 • 0 •• 0 • 0

• 0 • 0

0 • 0 •0 • 0 •

Figure 4.3.1 Planting patterns in mixed plots.Key: ., sprouted (i.e. multi or apical); 0, unsprouted (Cold).

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of 4 blocks. Spacing between rows was 76c~ ;illdbetweec ?luLts 36cm.

There were 3 tubers per r-epl i.c.s t ion, A~_l other details wer-e the same

as those described for Experiment F5.

4.3.6.2 Growth im:<lysm

All plan ts were h.u-vested 64 days ufter pLan ting Tnd growth para-

meters studied are presented in Table 4.4.8.2.

Snrout growth during storage

Ten tubers per treJtmclltwere used for sprout meusurement on 23rd

Earch, 8th and 14th Apr iL,

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L~.4 R~SUL'rS

4.4.1 Sprout growth durinG storage

The following dc,tes wer-e tiken as the days when dormancy of the

seed was broken : 27th December (Exp. F1) j14th December (Exp.F2);

15th December (EXIl.F3)j 22nd December (Expc, F5 and F6) when sprout

Length per tuber reached abou t 3mm (\1urr, 1978b), and the number of

si:.l.ydegrees shown 'tlerecounted from these dates. In the case of multi

t.r-eatmen t , it took about 50 - 60 day degrees to reach a sprout length

of 3mm tuber-1 (Figs.4.4.1.2 and 4.4.1.3) but day degrees shown here

are from the date they were moved to w~rm conditions (above 4°c) as

it was thought that tubers had broken dormancy by then. Residual

degrees of freedom and the \-ID.;; the ditn were anal.ysed is shown in

Table 4.4.1.1.

Total sprout length per tuber increased with increase in tuber

size (Figs.4.4.1.1; 4.4.1.2; 4.4.1.5). Tubers of the same size of

two var-Let i.ea: PentLmd CrowTl'...nd Record (Fig.4.4.1.1) had the same

sprout length. Linear regression between tuber size arid total sprout

length per tuber for both varieties (Exp. F1) of apical treatment

measured on 28th April (900 day degrees a':.Jo'le4°c from dormancy break)

accoun ted for 98)t.of the v.ir iance (Fig.L~.4.1.5). Similar relationships

are evident for other dates of measuremeLts (Fig.4.4.1.1). Sprout

numbers a.lao Lncr-eased with increase in tuber size ('l'ables4.4.1.2 and

4.4.1.3). Tot:...l sprout length per tuber Lncz-eaaed with increase in

day degrees above 4°c (Figs. 4.4.1.1; 4.4.1.2; 4.4.1.3; 4.4.1.6i 4.4.1.7).

Growth rate (extensior..)of the sprouts was increased by storing the

tubers in cold (3 ! 10C) before moving to warm conditions for sprouting

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/1

(Figs. 4.1+.1.1; !+.l~.1.2; 1~.1+.1.3; 1~.1'.1.7). For ex.unpl e in 1979 growth

r~te of Pentl~nd Crown from 11th Anri~ to 28th A~ril was 33.6mm per 100

day degrees above 4°C for mu.lti whi.Le this figure for'''picJ.l \-I3.S '7.66

(average of 3 Geed sizes). St.or i.ng the tuce r s ill cold C3 ~ 10C) before

moving them to \J,~.'.rm cor.di t ior.s for epr outir.g .i.lso il:cre;,.sed the number

of s or-out.s (T,:.;_bles4.4.1.2 c.lla i+.4.1.3).

In 1979 it vns observed fhu t some sor-cu ts s tcppe d growi.ng during

storace in the c~ie of 'ioical treatment. To invcstiCJ.te this in detail,

in the following two years, eyes were numbered (Cha?ter 4.3). If there

"Jere more th.in one snrou t on :.my eyCf then they 1derellso numbered, in

this vlay the growth of .ir.rl i vi duu.l sprouts wxs mo.ritored. In case of

ap i cal., on the bas i a of the length 0 f the sprouts :·t t'.J.irie:3.by the end

of the stor3.ge oeriod, sprouts were divided into four categories: those

tru thad r eache d be tween 2 to 3mrr:; 3 to 6mm; 6 to 9mm or over 9mm

(Fig.lt.4.1.L~J.). Sprout grmJt:l ra te of _ill apr out.e less th.m 9mmw;.rs

low but the .irnpor t.crt r-eauL t was that they stoooe d growing, while others

(>9mm) con t inue d to gr-ow (Fig.4.1~.1.1+). Simil:cr results wer-e obtained

for ~xperiments F3 ~nd F6, where for Gimolific~tion growth of only two

types of sprouts is shown i.e. <9md>3mm (Figs.LI.4.1.4 .md 4.4.1.7).

Fur the r it "k',S found th at apr ou ts whi ch con t inued to grm, were usually

present on the eye number 1 (rl';.cb1e1~.4.1.3). But if this eye was

damaged then it ','/:.1.S not necessarily so th.. t eye Humber 2 would continue

to grow, as in miny c.cses sprouts on tile he eI. end were seen growing

while others st opped , The length of the s or-ou ts 011 different eyes is

shown in Figure 4.4.1.8. In Expe r irne n t s F2 and F6 or.Ly u few tubers

had damaged eyes and so the s or-out Le.ig th is gr c.ite r on eye No.1, while

in Experiment F3 eyes vJere drmuge d HI m,::y tubers, ..nd so there Vias no

difference be tween different eyeoosi t ioria 'IS [,-,1",5 apr ou t length is

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';'2

concerned. Data in Figure 4.4.1.8 for growth of sprouts on different

eyes is presented only for 3 d~tes but similar results were obtained

for the remaining d~tes of me~surements also. Another factor which

should be tuken into cons i.der-at ion is the number of eyes present on the

tuber and data for this is shown in Table 4.4.1.5. Storing seed in the

cold (3 + 10C) before moving them to warm conditions for s?routing

removed the inhibiting effect of the dominating eye (T()ble 4.4.1.3 and

4.4.1.4). For aimplification, d.rta from different experiments is pre-

sen ted for the List measur-ement only but similar results were obtained

for other measurements also. Lowe r values for sprout length and their

number from e"J'eno ,? onwar-ds were not due to the fact that their growth

was inhibited by other growing sprouts but bec3.use these eyes were not

present on all the tubers (Table 4.4.1.5).

Total sprout length in case of mul ti vJ:.lsequal to api ca.l,by the end

of storage in all the four experiments (Figs.4.4.1.1; 4.4.1.2; 4.4.1.3;

4.4.1.7), but length of the ~ongest sprout (LS) in the case of multi

was much lower than that in the ap.ical in al.L the experiments (Table

4.4.1.6). To investigate this further, growth of the longest sprout of

2 types of treatments is presented in Figure 4.4.1.9. Although total

sprout growth rate waa much higher in multi (Figs.4.4.1.1j 4.4.1.2;

4.4.1.3; 4.4.1.7), due to intersprout competition, growth of the longest

sprout was not hi_gher than the apical.

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'73

Table 4.1+.1.1. Residuul. degrees of freedom (RDF) for vur i ous Figures

Fig./Table ~;o.

Fig. 4.4.1.1

Fig. 4.4.1.2apicalmulti

Fig. 4.4.1.3

Cipical (slow)ap i c a.l (fast)multi (slow)mul ti (fQst)

Fig. l~.4.1.4(a)

( b)

Fig. 4.4.1.5

Fig. 4.4.1.6

Fig. L~.4.1.7

For differen tcategories

iilldTables presented i~ this Ch~pter.

RDF

118

48

18

10

Remar-ks

Analyaed us comp.l eteLy random i.zeddesign(CRD) tuk i.ng tubers :_LS z-ep.Ldcutea,

sarne as abo veaame as above

Different number of tubers were involvedfor cornpu t ing their mean ;91 for first 8 dates u~d 46 for last date4534 for first 3 dates and 14 for last date20

Anu.l.yaed as split plot design taking seedsizes as main plots and categories assub-pl.o t.s, 'I'r-ays (8) were t.ake n as replicates.Anal ysed ilS CRD, trays (6) were taken asreplicates. (Fust not included).

4

Hean for 19 tubers.

For total sprout length meCln for 29 tubers.i r; cuse of ap ical,and 10 in case of mul ti

lm:Alysed :lS CRD, tuk i.ng trrys 0) as replicates.

continued ••

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Fig./Table No. RDF

Fig. 4.4.1.8(a) 66

(b) 22

cc)11050

Fig. L~.4.1.9(a)apical

multi(b)apicul

nul ti

Table 4.4.1.2 234

Table 4.'+.1.3

Table 1+.4.1.4 11

'I'able4.4.1.5

Tuble 4.4.1.6(a)

(b)

Rem~rks

Anulysed as split plot design, taking seedsize as ma.in plot and eyes as sub plot.'I'r ays (8) were tuken as replicates.CRD, tr'_;ys(3) tuke n as r-ep li.cctes,

All dates ano,lysed asFor first 2 datesFor Las t date

CRD LJ:ing trays asre?licates (fust notincluded ).

MeaD for 19 tubersNean for 10 tubersNean for 46 tubersHean for 14 tubers

AnalysedJ.s factorial (seed size X physiologicalage) CRD, Li.kbg tubers us re_?licates.

Mean for different number of tubers specified0.arlier for different experiments. In Exp.F3tubers from fast treatment were not included.

For Exp F3 and F4 an.a'lyaed as CRD bking trays(2) CiS replicates. In case of Exp , F6 theyare mean for 10 tubers.

Calculated from different number of tubers:Exp.F2, 70 for 62g tubers and 60 for 116gone;~Xl:).F3,125; Exp.F4, 19; Exp.F5, 40.

Analysed as f'ac torial (seed size X physiological:c..ge)GRD, taking tubers as renlicates.Diff. for different Exps.: Ex.?F2, 126; Exp.F3,121 (both ann.l.yae a as f'ac t.or-i.al, CRD). Exp.F4,me.:J.Ilfor 19 tubers; Ex.?F6, mean for 29 tubersin the cuse of api.cul,and 10 tubers in thec.ase of r.ml tie

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/5

Table 4.4.1.2. The effects of var-Le ty, tuber size ('rs) 'lid phys i.o.Logi cal,

age (PA) on s~rout number on 28.4.1979 (Exp. F1).

I'umbe r of apr ou ts 2mm tuber -1over

Pen tLand Crm-ID Fecord

PA Ap i ca l rlul t i ;":e:1.n PA Anic"l Nul. ti 11ean

TS,g 1'S,g

34 2.70 5.27 3.99 40 3.58 4.83 4.20

62 3.90 6.1(, 5.03 52 3.85 5.55 4.70

105 4.63 8.82 6.'72 63 3.90 6.30 5.10

mean 3.'74 6.75 mean 3.'78 5.56

P Crown Record

S2:D for age (mean ) 0.267 0.187

SED for tuber s i z.e (mean) 0.328 0.229

SED for body of the tdble 0.463 0.324

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Table 4.4.1.3. The effects of tuber size (rs) wld eye number on sprout

number of different size categor-y (SC), (2-3 days before

planting).

-1Number of sprouts, tuber on different eyes.

~xp. F3 Ex:). F4 Exp , F5 and F6.c:;xp. F2

53TS,g 62 116

Treatment 3.pic ,3-1 apic:;l multi(slow)

336(multi )

«o i.cal. 352 D(multi)

SC, mm 2-9 >9 2-9 >9 !2-9 >9 2-3 >3 2-3 >3 '2-9 >9 2-3 >3

Eye no.

1 0.13 0.58 0.08 0.7 0.39 0.30 0 1.21 0.16 1.63 0.14 0.52 0.2 1.0

2 0.23 0.08 0.35 0.03 0.39 0.17 O.O,?0.71 0 1.75 0.38 0.14 0.2 0.8

3 0.18 0.10 0.3 0.10 0.28 0.35 0.09 o. '/9 0 1.3'?0.34 0.10 0.3 0.8

4 0.15 0.08 0.48 0.08 0.35 0.35 0.29 1.07 0.05 1.32 0.45 0.14 0.2 0.7

5 0.25 0.08 0.33 0.13 0.46 0.26 0.29 1.61l- 0.05 1.32 0.45 0.14 0.1 1.0

6 0.10 0 0.25 0.13 0.26 0.24 0.14 1.36 0.21 0.95 0.38 0.03 0.4 0.7

7 0.13 0.10 0.13 0.05 0.41 0.20 0.36 0.'110.42 1.32 0.10 0 0.3 0.9

8 0.15 0.10 0.10 0 0.26 0.04 0.21 0.57 0.26 0.84 0.17 0 0.4 0.3

9 0.10 0.25 0.10 0.13 0.13 0.24 0.07 0.21 0.16 0.63 0.14 0.06 0 0.1

10 0.15 0.13 0.2 0.08 0.02 0 0 0.14 0.05 0.26 0.14 0.03 0.1

11 0.05 0.08 0.05 0.10 - - O.O,?0.14 0.05 0.16 0.0'70.03 --12 - 0.03 0.03 0 0 0.05

Total 1.6 1.45 2.4 1.53 2.96 2.02 1.5 8.57 1.42 11.57 2.75 1.34 2.2 6.3

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Table 4.4.1.4. The effects of tuber Slze and eye number on total sprout

length (2-3 d~ys before plonting).

Snrout length, mm tuber -1 differenton eyes

Experiment F3 F4 F6 F6 F6

Treatment i-1ulti B36 352D 280D 232D(slO'.-:) (multi )

Tuber size,s '78 204 53 53 53

Eye No ,

1 11.42 19.12 9.1+5 9.25 9.15

2 6.91 19.14 6.40 6.70 8.25

3 6.54 16.72 6.15 6.75 5.45

4 11.61 11~.49 5.15 3.85 9.35

5 13.98 14.53 7.77 5.75 6.05

.6 11.96 9.75 7.30 6.80 3.15

( 6.82 12.42 6.30 6.10 2.95

8 4.86 7.34 2.40 3.90 3.55

9 2.14 5.68 0.55 2.35 0.50

10 1.18 2.72 1.00 0.20 0.60

11 1.20 2.01

SEB 1~65 3.01

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'-j I)(U

Table 4.4.1.5. 'I'he effect of tuber size on number of eyes.

Percent~ge of tubers had these eyes

Lxperiment F2 F3 1"4 F5

Tuber size,g 62 116 '('8 204 53Eye .No.

6 97 98 94 9? 100

7 94 98 82 93 100

8 86 95 50 90 97

9 61 88 29 80 90

10 19 65 14 60 62

11 6 23 8 37 24

12 0 12 0 17 3.4

N.B. At least five eyes were present on every tuber.

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Table 4.4.1.6(a). 'I'he effects of phye io.Logi ca.l age (PA), var i e ty and

tuber size ('rs) on .leng th of the longest sprout,

-1 ~ 4 )mmtuber on 20•• 1979 (Exp.F1 •

:?entl::md Crown Record

PA Anice,l l';ul ti l'iecm 0' • ' 1 j,lul ti Mean'11 i~PlC:,L

T.3,g TS,g

34 19.05 8.68 13.86 40 16.35 10.50 13.43

62 18.70 9.07 13.89 52 17.75 11.22 14.49

103 20.43 9.85 15.14 63 19.32 10.68 15.00

mean 19.39 9.20 me.in 1'7.81 10.81

P Crown Record

SZD for age (mean) 0.440 0.415

SED for tuber size (me an ) 0.538 0.509

SED for body of the t.ab l,e 0.761 0.720

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Table 4.4.1.6(b). The effects of physiological age (PA), sprouting

treatment (ST) and tuber size (TS) on length of the-1longest sprout, nun tuber (measured 2-3 days before

planting).

Exp. F2 li'3 li'h F6

TR SPL TR SPL 'ill SPL 'l'R SPL(mm) (mm) (mm) (mm)

TS,g ST

62 14.? fust 29.8 1336 16.0 + 0.48 920D 19.6 ~ 0.56

352D 10.9 ~ 0.87

116 18.5 24.4 280D 10.5 +slow - 0.55

SED 0.53 SED 1.0'/ 232D 10.4 + 0.50-

PA PA 184D 9.1 ~ 0.63

apical 22.3 aDicul 31.6 128D 8.3 + 0.62-rr:ulti SOD 4.9 +multi 7.0 15.3 - 0.53

0.54 SED 1.20 48D 1.77 +SED _ 0.31

\.JherejTR = treatment; SPL = sprout length.

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GO

rlI

IQ

50_1 /II

: I~.O il 1!P'I 1/

-',o. I rp~j '1 1/1

I / / I .') r-. r!J/ IGU_ AI

10!

Pentland Crown ISEDso l05gL::.. 62go 34g

ApicalMulti

I

i-I () ~-------r'-'--"-"'-- T----·-··---T··------···T ...····-,··---···-·----·---r--·----·-····--r----,

100 20 0 J00 (~0 0 ~)(; i ) C()C) 'H):') t~C0 900

,,

iC (J' I.J. -i

I,;i ffJI40_! Iff

IIII Cj)

30.1 /1 I1/1 IIII

20_ I{~~

Recordo 63gL::.. 52go 40g

ApicalMulti

ISEDs

100-1 O-+----,---·--···r··-·-- ----T- ..---.T-----.-.-.--.-T.--,----------,-.---,

200 300 400 500 GOO 100 800 900oDay degrees above 4 C from dormancy break

Figure 4.4.1.1 The effects of tuber size, variety andsprouting technique on total sprout length duringstorage (Experiment Fl).

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75

GO.

i.5

30.

15

rlI

~..c:to 1:5j;:l(])rl

+';:joaGO.Ul

rlcd+'o8i.5

30

15

1;'rilll~

./1SEefor mean

/.I

o Slowo Fast

ApicalMulti

/ I

// I

/ ./8........--:r//

'/

O-+----,----r----.-50 150 250 350

'-'550

1

950i.50 G50 750 850Figure 4.4.1.3 The effects of sprouting technique on total sprout

length during storage (Experiment F3).

6. llfigo 62g

ApicalMulti

II SEe

I

I

I

-- ..--,----,---- ---T------,---- 'T 1 --·--1

150 250 350 oi.50 550 GSO "lSC, 850 950Day degrees above 4 C from dormancy break

Figure 4.4.1.2 The effects of tuber size and~routing techniqueon total sprout length during storage (Experiment F2).

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rlI

50

40

30

20

10

Experiment F3 I SEDsI ~/

I .0./

./

I /.0/

.ff»:_./

I B-:-:

.ff»:_./

.Er»:/

B-:::r:. ././ 1---8----9--- __Df------f.=Of------<Ot:t---0

~-o

o >9mm

o <9mm

I

II

~ 0-+---,---.----.--,-----.---.---.-----.-----,'§ 50 150 250 350 450 550 550 '150 850 950+'

gj,.; 50+'tlOs::OJrl

+'g 40HP,U)

30 I

Experiment F2

o >9mmA 6-9mm

o 3-6mm

+<3mm J: SEDs.a

./_-8'

:l: .a-r _

.a--r E-_/

_/

20

r_-Er

2-10

-A------i:r-----/::;-- -ubA----A----8----t:r-----:

oj~~~~t;~~-~-- -?:=~=_u~------ J T-- _~====~====~====~c==~

50 150 250 350 ~50 550 550 '150 850 950Day degrees above 4°c from dormancy break

Figure 4.4.1.4 Growth of sprouts of different categories(see text for details) during storage.

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1;~CiIII

'1()~:4I

••,;613')1'-'lII

10"'1'--"-, --T----T-···- . l" ....- 'I

50 150 "'J'Day degrees above 4°c from

dormancy breakFigure 4.4.1.6 Total sprout length

per tuber during storage(Experiment F4).

rlI

J.,(!).0::l+-'

~".£1

+-'~~(!)rl

+-'::l0J.,P;Cl)

~)s_,I

III!

rl '51I (. 1,J.,

I(!).0::l+-'

~ r55 I"-I" i

.£1,

+-'

IQo~(!)rl

25~+-'::l0 IIJ.,

~

1Sj.-,

III51100

~ma for meanI rl

1

H(!)

.g+'

~ 50",.s::::

+'till 0~(!) £Irl /

+' 40 /;j .00 /HP<Cl) tD

30 ---I30 50 70 90 no

tuber Slze, g

I

I

Figure 4.4.1.5 The relationship betweentuber size and total sprout lengthmeasured on 28 April (Experiment Fl)

y=20.48(+1.38) + 0.36(+0.02)x% varian~e accounted f~r 98.2,residualstandard deviation 1.23.

I SEa for meanI Total

SEDs'>9mm

~/

.R!:./

///

;{IISEa' for mean

<.9mm

Figure 4.1~.1.7 Total sprout length and sprout length of differentcategories during storage. Experiments: F5 and F6.Key: __ , Apical; --, Multi.

I

'1'-- 1 1 r-·-·-----T-----T ..'-----"---'--'---,

200 30Cl C; IJU ~-)ClU h:i~' 'N)U t3 00 80 ()Day degrees above 4oCfrom dormancy break

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Experiment F372 520 936o 0 t:::.

*

3_

oJ--,----------r------r---r---- --r-------T-----T-------r--------r-- ----,-----1o 1 2 3 4 5 5 1 8 3 10 11 12

....c: 0+' " . --r"eoI=l r' '/'1 r;> J I. 5 5 9 10 11 12(!J ur-I

+' -15_ Experiment F5::s I0

1,,11-<P< *Cl)

:J112 568 9040 0 t:::.

I

I Ii I,

5JII

II

3~

01 -T T- , - -- - I - r- 1

0 1 () J ~ 5 5 '1 8 9 11G

Eye Number

r-II

Experiment F2

56 416 952o 0 t:::.

*

I I

'.. Figure 4.4.1.8 Total sprout length contributed by differenteyes during storage, for apical treatment only.* Day degrees above 4°c from dormancy break and SEDs to

cQmpare between eyes.

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30_

24

18

12

8 :c

Experiment F1I SEs for mean

o Apical (slow)o Multi (slow) I

I

I

.I

IIjZj

~/O~-----'------'------r---------- r-------,-----,-------r----,50 150 250 350 450 550 850 '750 850 950

~.r:: 30_, F6+' Experiments: F5;bOc IQ) Irl

I

+' I o Apical;::!0 24J o Multi~c, ICl) i

I I SEs for meanII

18_

I

12_ .r II

8

O-,---------I-----------r------ ----r---- -----r--" ---I' - - 1--- - --------'1------- - -,-------,

50150 250 350 450 ~<50 850 '750 850 950oDay degrees above 4 C from dormancy break

,;

Figure 4.4.1.9 Length of the longest sprout during storage.

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4.4.2 Experiment F1.

Soil temperature, at 10cm. depth, screen, max and minimum air

temperature and rainfall data are presented in Figures 3.3.3a, 3.3.2a,

4.4.2.1 .emergence ar:dstem number

The variety Record started emerging about 10 days after Pentland

Crown, but emergence was more homogenous in this variety thus the

difference in time to 501b emergence was reduced to 8 days (Fig.4.4.2.1.).

Once emergence had started the rate of emergence was higher in Record

while physiological age and spacing did not affect it. Inspite of

differences in the length of the longest sprout at the time of planting,

physiological age did not affect the emergence, may be due to the fact

that total sprout growth rate was higher in case of multi (Fig.4.4.1.1.).

Stem number stopped increasing after the end of June and thus were

averaged for the various dates of gr-owth analyses carried out after that

as there wetsno significant difference between different dates. The

total number of stems per unit area were higher at 30cm. (22.11,M-2)-2spacing as compared to 4Ccm. (17.09,H ) but this increase was in pro-

portion to the increase in number of plants per unit area. Apical

treatment increased the numb=r- of br-anch stems in both varieties

(Fig.4.4.2.1.). Record had a higher number of branch stems but a lower

number of main stems and total stem number was also higher in this

variety (Fig.4.4.2.1.). There \vilS no increase in the number of axillary

branches (AB) after the end of June and so they were averaged for all

dates of gr-owth analyses. AB wer-e higher in Pentland Crown (Fig.4.4.2.2.).

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Apical treatment increased tl:e number of AB in Pentland Crown but

decreased in Record (Fig.4.4.2.2.). It may be explained by increase

in total stem number by this treatment (ApicaJ.) in Record, which in-

creased shading of the lateral buds. Similarly planting at closer

spacing also decreased the number of AB. In Pentlund Crown about 98%of the AB present were contributed by the ma.i,nstems, this may be

because branch stems were fewer in number (Fig.4.4.2.1.) and further

they were smaller in size (Fig.4.4.2.15), thus lateral buds may have

been shaded and stayed dormant. Development of AB depends on the

release of dominance from the apical bud, which may be related to the

lower auxin production by the lateral buds. Thus development of AB

may be related to the speed of ground cover by the crop canopy. To

investigate this the le~f area duration for a period of 30 days from

the date of 50}~ emergence (LAD'E30) was calculated and plotted against

number of AB present (Fig.4.4.2.2. ). Number of AB present decreased

with increase in LAD'E30 i.e. with the increase in speed, with which

ground was covered by the canopy. Speed of ground cover was faster in

Record.

4.4.2.2 Stolon grO\vth and development

The variety Pentland Crown increased stolon weight (Fig.4.4.2.3)

for a longer time and this may be related to delay in tuber initiation

(TI). The variety Record initiated tubers 5 days before Pentland Crown

(Table 4.4.2.1), when counted from the date of 5O;~ emergence. Thus while

Record started feeding its tubers, assimilates in Pentland Crown may

still have been used for stolon growth. Stolon growth slowed down

after TI in both the varieties (Fig.4.4.2.3). Early in the growing

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season before TI, apical treatment increased stolon weight per unit

area (Fig.4.4.2.3). Percentage of stolon dry weight out of total dry

weight was also higher in this treatment. Spacing, did not affect

stolon growth or its number (Fig.4.4.2.3). StoLon number were also

unaffected by 9hysiological age. Like stolon weight, stolon number

also stopped increasing after 'I'L, Stolon number were higher in Pentland

Crown. In general before TI stolon gr-owth \Vus found to be linearly

related to the LAI of the canopy (Fig.4.4.2.4).

4.4.2.3 Growth and development of stem and leaf

In general main sternswere slightly longer than the branch stems

but on an overall basis they were longer in Pentland Crown than in

Record (Fig.4.4.2.5). Physiological age mld spacing did not affect

the plant height. Total number of stems were higher in Record (Fig.4.4.2.1)

and they had higher weight of under ground stem (UGS) (Fig.4.4.2.7).

Increase in weight of above ground stem (AGS) was not proportional to

the UGS, in fact Record had decreased AGS wei.ght , This may be explained

by the decrease ir-height of the plant (Fig.4.4.2.5). Physiological

age did not affect the stem weight, while plonting at closer spacing

increased stem weight (UGS as well as AGS) per un i t area (Figs.4.4.2.6 and

4.4.2.7). Thi& may be due to higher number of plants per unit area.

Record emerged 8 days later (Fig.4.4.2.1) and thus had lower LAI

early in the season (Fig.4.4.2.8), when considered from the date of

planting but it was much higher in Record when considered from the date

of 50;6 emer-gerice, and the rate of increase in LAI was also higher thus

later on it had higher LAI than Pentland Crown, but later it declined

rapidly due to earlier senescence (Table 4.4.2.1). Apical treatment

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decreased LAl (may be due to decrease in stem no.) Although the effect

was not significant at any date of Growth analysis but was consistent

during most of the growing season (Fig.4.4.2.8). Planting at closer

spacing (30cm.) increased the LAI and the eI'f'ecttwus consistent through-

out the growing seaSOL (Fig.4.4.2.8). This may be due to the higher

number of plants per uni t area. Overall about SO;:{, of the LAI was

contributed by the rnain stems in Pentland Crown while in Record this

figure was 52%. Apical treatment increased the number of branch stems

thus decreased the proportion of LAl contributed by the main stems,

from 82 to 57% and this degree ot:decrease (25%) was the S3ll1efor both

varieties. Although there was no increase in the number of AB after

end of June but existing branches continued to grow. Size of AB may

be studied by working out their leaf area sep&rately and is presented

in Figure 4.4.2.9. LAl contributed by the AB (LAP AB) was higher in

Pentland Crown. Number of AB were also higher in this variety. Closer

spacing and multi treatment decreased the LAI'AB. This may firstly be

due to a lower no. of AB present in these treatments and secondly due

to higher competition, as they had higher number of stems (main + branch)

(Fig.4.4.2.1). Higher LAl' AB in Pentland Crown and the treatment apical

was mainly due to the higher number of leaves present on the AB of

these treatments. There vJaS also a slight increase in the average leaf

size of the leaves present on the AB, due to these treatments.

Apart from the leaves present on the AB, the total leaf numbers

were ned.ther affected by the varieties nor by the physiological ages.

\llhileplanting at closer spacing did significantly increase the total

leaf number (Fig.4.4.2.11) but average leaf size was slightly decreased.

Although total leaf numbers were not affected by the varieties and

physiological ages but the important result was that leaves coming from

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different type of stems were affected. In the case of Record more

leaves were contributed by the branch stems and the reverse was the

case for Pentland Crown (Fig.4.4.2.12). Apical treatment which in-

creased the number of branch stems, also increased the proportion of

leaves contributed by the branch stems (Fig.4.4.2.12). Leaves present

on the ABwere much smaller in size, thus average leaf size for main

and branch stem presented (Fig.4.4.2.13) was calculated w.i thout

including these leaves. Leaves coming from the branch stems wer-e much

emal.Ler and difference \'las gr-ea te r in the variety PentLand Crown

(Fig.4.4.2.13). Branch stems of multi treatment had much smaller leaves

than those of the api ca.l treatment.

Decrease in leaf size resulted in di f fer-en t ial, vu.l.ues for leaf to

stem ratios (Fig.4.4.2.14). Br-anch stems had a lower Leaf to stem

r a t i,o and the difference was gre ..rt er in Pentland Crown, Leaf to stem

ratio of br-anch stems W2S increased by the apical t.r-ea tment

(Fig.4.4.2.14). Spac ing did notJ.ffect the leaf to stem r at i o of any

particular type of stem. In gener:li leaf to stem r-at i,o vias slightly

decreased by planting at closer spacing.

Total above ground stems are usual.Ly accep ted as criteria for plant

population in po tato , thus it may be important to study the size of the

different type of stems. Size of the stem is presented in term of LA

per stem (Fig.4.4.2.15). Branch stems were much smaller than the

corresponding mai.n stems .md this difference was greater in Pentland

Crown. Apical treatment increased the size of muin stem in Pentland

Crown but decreased it in Record. However size of branch stem was

increased by the api.ca.L t re a tment in both varieties but degree of in-

'. crease was more in Pentland Crown (Fig.4.4.2.15).

The effects of varieties and spacings on specific leaf area is

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86

presented in Figure 4.4.2.10. Specific leaf area was higher in Record

and at closer spacing, this may be related to the mutual shading by theleaves as both these treatments increased the LAI also. Decrease in

specific leilf area due to decrease in irradiance was also found in the

experiment GR2 (Chapter 2). Physiological age did not affect the

specific leaf area.

4.4.2.4 Light Interception

Figure 4.4.2.16 shows the proportion of PAR intercepted by the

crop canopy. Quantity of PAR intercepted by the canopy viaS increased

by planting at closer spacing and this increase was in proportion to

increase in the LAI (Fig.4.4.2.8). Effects of variety and physiological

age on PAR interception were also similar to their effects on the LAI

(Figs.4.4.2.8 and 4.4.2.16). Helationshin between LAI and light

interception is shown in Chapter 4.4.6.

4.4.2.5 Tuber growth ~~d development

'l'uberinitiation (TI) was considered as the date when tuber dry-1we igh t of 0.25g plant was reached and was calculated by interpolation

from the growth analyses and addition:u sampling data, taken frequently

during the early growth of the crop. TI in Record occurred 3 days later

than in Pentland Crown (Table 4.4.2.1) but when considered from the date

of 5CJl;6emergence it was in fact 5 days ear-Li er- Ln Record (Table 4.4.2.1).

Apical treatmeEt enhance, TI (one day) by enhancing the emergence

(Fig.4.4.2.1) thus there was no difference in the number of days taken

.for tuber to ini tiate when coun ted from the date of 5a:!~emer'gence,. ,

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Table 4.4.2.1. The effects of variety, physiologic~l age and spacing

on number of days taken: from planting to tuber initiation eTI) (TIP);

from 50}b emergence to TI (TIY); from planting to senescence (S£1JP).

'rIP TI.t: SL'NP

V.J.riety

Pentland Crown 49.92 20.83 159.8

Record 52.67 15.58 137.9

Physiological age

lvIulti 51.92 18.25 150.3

Apical 50.67 18.17 147.5

Sp~cing

30cm 51.50 18.17 149.3

40cm 51.08 18.25 148.4

SED 0.453 0.497 2.90

Record gave consistently higher tuber weights as well as tuber numbers

throughout the growing season (Figs.4.4.2.17 and 4.4.2.18) but final

tuber yield was higher in Pentland Crown because it stayed green in the

field for a further period of 22 days after Record had senesced

(Table 4.4.2.1). Just after TI apical treatment did give higher tuber

weight and number (Figs.4.4.2.17 and 4.4.2.18) but this difference soon

disappeared and final tuber yield was not affected and tuber numbers

were increased by the multi treatment. Planting at closer spacing'increased tuber number us well ·"stuber we igh t (F;g 4 4 2 17 d 4 4 2 18)~ • .L s. • •• a..'1 ••• •

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8b

The effect of sp::..cingon tuber number \'Id.S more, early in the se3.son

(Fig.4.4.2.18), which may be attributed to the incre:J.sein LAI by plant-

ing at closer sp:J.cingbut subsequently percentage of increase in LAI

decreased along with the season due to higher interstem competition,

thus differences at final harvesting were reduced. Tuber numbers are

usually related to the total stem numbers as the latter increase the

LAI at the time of TI and this was found to be the case in the present

experiment (Fig.4.4.2.19). 'I'ubernumbers which result seems to depend

on a~similates available for their growth for some period after TI.

Thus it was of interest to look at this in further detail. Leaf area

duration for a period of 30 days (LAD'TI30) was calculated from the date

of TI and is plotted against tuber number (Fig.4.4.2.20). About 98.7~

of the variance in tuber number was accounted by linear regression

between tuber number and LAD'TI30 in Record and Tc~J in Pentland Crown.

Record and Pentland Cr-own had two separ-ate significantly different

regression lines. Record developed 1.10 tubers per unit of LAD'TI30,

wh iLe this figure for Pentland Crown was 0.78. This difference may be

explained by the fact that a higher perceZltage of assimilates were

allocated to the tubers in the case of Record (Fig.4.4.2.25).

Total tuber yield was found to be linearly related to the total

leaf area duration (LAD) accumulated over the whole season by the crop

canopy (Fig.4•.4.2.21). Tuber yield per unit of LAD \'I<.3.S 5.31 in case

of Pentland Crown and 6.34 in case of Record. Higher efficiency of

Record may be related to the higher percentage of assimilates allocated

to the tubers (Fig.4.4.2.25). Since higher tuber numbers were found in

Record, assimilates available for the growth of individual tuber were

lower, which resulted illhigher proportion of medium sized tubers.(Fig.4.4.2.22). At final harvesti.ngRecord gave 7Z;b of the medium

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sized tubers <32-51mm), while this figure for Pentland Crown was only

3716most others being larger. Closer spacing (30cm) and multi treat-

ment also increased the proportion of medlum sized tubers but difference

was not significaDt. Tuber size ~ay be related to the yresence of AB.

Because AB present, may increase the amount of aas imiLates ava.i.Lab'Le

for a particular tuber to Grow .md thus increase the tuber size. To

examine this Le af ar-ea duration contributed by the Ai3 (LAD'.AB) '!las cal-

culated and lS plotted og,Clinst tuber weigh t over 57mm (Fig.4.4.2.23).

Lme ar regression between tuber wei.ght over 57mmand LAD'AB accounted for

631b of the var-Lance in the tuber weight over 57mm.

4.4.2.6 Total dry matter accumulation

Roots were not collected and those present on stems and stolons

were removed and discarded, thus the tobl dry weight (TD~J) presented

is excluding roots. Later in the season (end of July) leaves started

to falloff and were not collected from the ground. Pentland Crown

completed 50% emergence about 8 days earlier than the variety Record

(Fig.4.4.2.1) and thus had higher total dry weight early in the season

but crop growth rate was higher in the variety Record and thus TDlv

produced by the canopy of this variety was higher during the middle

part of the gr~:)\oJingseason CFtg.4.4.2.21+). Later it senesced before

Pentland Cr-own (Table 4.4.2.1). Planting at closer spacing also in-

creased the Tl;r.{per unit area (Fig. 4. 4. 2. 24) whi ch may be due to the

higher number of photons intercepted by the canopy (Fig.4.4.2.16).

Physiological age did not affect the TD;.,r(Fig.4.4.2.24). Various

components making up the TD'\oJ may be of importance as tubers are the

economic part of the crop. Record allocated a higher percentage of

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')()

assimilates to the tubers (Fig.4.4.2.25) and less to the AGS. Apical

treatment also resulted in an allocation of a higher percentage of

assimilates to the tubers ana this effect was consistent throughout the

growing season (Fig.4.4.2.25). Planting at closer spacing increased

the percentage of assimilates culocated to the stems and decreased to

the tubers.

4.4.3. Experiments: F2j F3j F4.

4.4.3.1 Emergence and stem number

As in experiment F1 (1979), apical and multi treatments of

experiments F2 and F3 started emerging at the same time (28 days after

planting (DAP) (Figs.4.4.3.1 and 4.4.3.2) but the time to reach 5~fo

emergence in case of api caL was 34.3 days in experimen t F2 and 31.7 in

experiment F3, which may be due to the fact that apical in experiment

F2 made more total stem number and proportion of branch stems was

higher (Fig.4.~.3.2 and Table 4.4.3.1) and branch stems may have emerged

later. Cold started emerging 34 DAP but emergence in cold was more

homogenous and difference in time to reach 50% emergence was reduced

to 5 days compared to multi and 3.7 days compared to apical

(Fig.4.4.3.1), while in experiment F4 the difference in time to 50%

emergence was 8 days, between S25 (cold) and B.36 (multi). This may

.be explained by the difference in seed size in two experiments.

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"~Ct-io 30~ • •!-<Q)

~z20.

25

20

15C\JI

S"!-< 10

Q)

~$:1

s 5Q)

+'Cl)

50.

Stem number averagedoVe!' Va.r10U~ growthanalyses.

SEDsI II ~ __----{J

I ,r ~~~,..¢' .... ~

1'/ ,,~1

o

•• o

o

16 20

•0 •

1 II.ijI I 0I II I 0~ Date of 50% emergence, II I anditsSEDI.I"

•5 10 15 20 25June

25 30May

Figure 4.4.2.1 The effects of variety and physiological age onemergence.

o o y =78.24 (2:_11.39)-2.24(2:_0.53)x% variance accounted for 70.4residual standard deviation = a 64.

10·ur-----------r----------.----- ,- ~20.0

LAD/E30, daysFigure 4.4.2.2 The relationship between leaf area duration,

accumulated over a period of 30 days from the date of 50%emergence (LAD/E30) and the number of axillary branches (AB).

10.0 15.0 30.0 30.0

Key.:,0, Pentland Crown; 0, Record; open symbols, apical; closedsymbols, multi.

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Variety

o Pentland Crowno Record

7.0 200C\J I I III s

b.O 5.6 I SEDs 17.-+'..c:

~ ~eo'r-! r \Q) I \~ 4.2 I \ C\J 15» h--- ---Q._

IS IH SEDs.-0 "D .- ~Hs::: Q) ,0

~ 13 1r-1 2.80I+' s:::

U) \s:::

~0 \r-1

1.4 $112\U)

\0

0 9

Physiological age Spacing7.0 7.0

1.4

C\JI

S

b.O 5.6 5.6.-

+'~.r-!Q)

~ 4.2~.-0

s:::~ 2.8o+'U)

4.2

o Multio Apical

2.80 30cm0 .40 cm

1.4

ci

0Sept. June July Aug. Sept.

oJune July Aug.

Figure 4.4.2.3 The effects of variety, physiological ageand spacing on stolon dry weight and the effects of varietyon stplon number.

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2.50 y == 0.067 (~ 0.279) + 2.49 (~ 0.51) x

% varlance accounted for 73.7Residual standard deviation = 0.29

2.10

0

C\I 1.70 0ISeo

+'..c::eo 1.30'rl 0OJ~

~'d

s:: .900 0rl0

0 0+'co.50 .20 .40 .6 0 1.00LAI

Figure 4.4.2.4 The relationship between stolon dry weightand LAI. Key: o , Pentland Crown; 0 , Record.

"+'fib'rl 30OJ..c:~~ 20p.

OJeo~ 10~

60SEDs

50

40

_......0-_ 0

_,.O'" -0- - - -/ -0/

o//

......a'o 0

--0

Pentland CrownRecord

o

o

-Iune SeptemberJuly August

Figure 4.4.2.5plant height.

The effects of variety on average

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oOO-L------.---------.------------r-----

X102085

042C\J1

Gb 021~Cl)Cj«

000

X102 Spacing085

064

042

021

064

042

064

021

Variety

II

o Pentland CrownORecord

Physiological age

./..8- ___

Er./»:

ra-j

jOMultiOApical

____-El

030cm040cm

June July~--

August Sept.

Figure 4.4.2.6 The effects of variety, physiological age andspacing on dry weight of above ground stems (AGS).

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1 I I I-: ---s- - -B- I

Id- --- -g - "El I1// ~~(Zl

I./d

Variety25

I

I20

15

10

o..!- "I _

25 Physiological age

20

15C\J 10I

6h..r:fl 5()~

0

25 Spacing

20

15

10

5

o~ ~ __June July

Figure 4.4.2.7

ISEDs

o Pentland Crowno Record

------.----------------------------- T----- --------

o Multio Apical

T------ ------------ --1

---- B"13-- -B----£J

o 30cmo 40cm

August Sept.

The effects of variety, physiological age andspaclng on dry weight of under ground stems (UGS).

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2050Variety

I I I I II~~2000 I 13' Dj

-Er \ ISEDsy:r-1050 I \

\1000 ~

050~

0 Pentland Crowno Record

000

2050lPhysiological age

2DOO~

1050

1000

/050H

:§ 000

2050 Spacing

2000

1000

050

o Multio Apical

o 30cmo 40cm

JuneoOO-'-----.-----------r------------r-----

July August Sept.

Figure 4.4.2.8 The effects of variety, physiological age andspacing on leaf area index (LAI).

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Varietyo SOl o Pentland Crown

050 Spacings

040 0 30cm0 40cm

030

0201

010

040

030

020

010

o SOl040

030

0201~ Di°l<,H

j 000

I II ISEDs

o Record

~--------~

Physiological age

o Multio Apical

)cl

/ "" _-f)/ a--.0Id-_ .// -s. ./

/ -s/ --~-~o-

·~-----=--'---~----·-~--r

Q/ \/ \

.0/Jd-_

/ -s../ ~~~~---~

/.0000~ ~ -. ~ ~ __

Figure 4.4.2.9

June July August Sept.

The effects of variety, physiological age andspaclng on leaf area index contributed by the axillarybranches (LAIjAB).

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2050Variety

I SEDs2025 I T r I

:r :r

2000

1015rlI

bD 1050_C\],a

1025~jP-.U)

20 50~

·2025I

2000~

1015

11050j

1025

o Pentland Crowno Record

Spacing

o 30cmo 40cm

June July SeptemberAugustFigure 4.4.2.10

280l Y'I I III <, ~ I

l ~ '~

21011 J. ./'-B- -P ..._B- __ £ 8- ___

1[,0 I _0' - i3- - -8. <,

/ R~o 30cm ~'f.)

C\JI

70

The effects of variety and spacing on specificleaf area (SPLA).

I I IISEDS

o 40cm

0-'- -.._June

-----------.--------------rJuly August September

Figure 4.4.2.11 The effects of spacing on leaf number,excluding the leaves coming from the axillary branches.

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2-10,I!

i-158!

1

!i!I125_:

II

8~_~

C\JI

"HQ)

§2-10__>=1

Ict--l ,

ItU IQ)

H I

158_~I

iiI

125JI

Variety

IIII o Record

o Pentland Crown

II SEDs0J./ \

)d/ \ / <,

/ \ / ':/J M 'B- - - -R ""-

I

-------1----------------------------y-----

Physiological age o Multio Apical

84

1//~" /

.G..~

'42 I e' <,

B_~I-e--

° -G:- -==- ~--June

Figure 4.4.2.12

,-------------------,-- h -------------- IJuly August Sept.

The effects of variety (V) and physiological age

(PA) on leaf number contributed by the differant types of stems,excluding the leaves coming from the axillary branches. SED onthe right side is to be used for comparing means within samelevel;:;of V or PA. Residual degrees of freedom is41.Key: ---, main stem; ----, branch stem.

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1025

1000.

II

o 15~I

0501

III025

Variety

SEDsIIIIII

.0/

/./ B-- --..-0

-- e- ./

o 00-"-------·--,---·---------···---1---··--- .---.--------..-----

..s 102 5l

Physiological age.,.,Ul

IHtilQ)H

1000

050

025

000Sept.

Figure 4.4.2.13 The effects of variety (V) and physiologicalage (PA) on leaf size (leaves coming from the axillarybranches were not included). For meaning of SEDs, symbolsand residual degrees of freedom see figure 4.4.2.12.

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,

I

20 10~

I104°1

i

01°1o ooJ_--~~~---'-T~~----'----~----'~-----~--'---T~'--"-~-~--------

3 050_Ii

iII

2 a 801

0'r-!+'ajH

S 3050_,OJ+'en i0 I+' I'\-i Iaj

2 0 80~OJH

II

2010JIi,iI

1040J

i

010

000

-,-,

'M- --- .";' \

\\ ..0

'13- .--- -- B- -- --e- ./

Variety

I III II

Physiological age

IIII II SEDsB - --H- - -:-

//

........... -=- - - -e- ---'& - "'"

..e---8---B---

-----~----------T· ~--._._-_.._._...._-

June July

Figure 4.4.2.14.

I ~--r--Sept.August

The effects of variety (V) and physiologicalage (PA) on leaf to stem ratio (dry weight basis). Formeaning of SEDs, symbols and residual degrees of freedomsee figure 4.4.2.12.

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Pentland Crown

30 Multi

24

18

~I

~+'Ul

Multi

18_

12

6

O-'---r-----,----------,--JuJ.,y Aug.

Apical

Record30,

I Apical

24.

18

I II(2)

0._-.-July Aug.

Figure 4.4.2.15 The effects of variety (V) and physiologicalage (PA) on stem size.Key:_, main stem;for comparing means within same levels of V or PA.

branch stem. SEDs (2) is to be used

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Variety I80 I

I I IB-

/ I I'10 I I_-8'- \.

f Q_ \1 I I60 I p- \/ tJ SEDs

~30.

r / 0 Pentland Crown40 I 0 Record

<. y3D

80

10.

60

50

{.o

Physiological age

o Multio Apical

3 0 _~ ~

80

'10

60

50

{.o

30_ rj

. r------ --_ ... _-_ ...._---,- ----~-- - .... -- 1

Spacing

p./

.0--

oo

/-----..~·---..-----I------ ..-----~----.--

June July Sept.August

Figure 4.4.2.16 The effects of variety, physiological age andspacing on photosynthetically active radiation (PAR) interception.

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1050 Variety

eos:1'rl+>(I)

ria;ro :>s:1 H

I'rl ror:r..::r::rSEDs0

II 0

B'-'...---

0 Pentland Crown0 Record

8{'0

{'20

630

210

1050_ Physiological age-)

-El

8{'0 B./

.Er630 -:

C\J ../I

Gb .>{'20 -+>..c: /'eo 210 if''rla; ,/ 0 Multi~~ .s 0 ApicalH ~'d 0 /'

Ha;..0;:l8 1050

l Spacing0

8{,OJ -El./' 0

I»>

630 '7.0

/'

{'20/'210 -: 0 30cm

0 0 40cm

June July August Sept.

Figure 4.4.2.17 The effects of variety, physiological age andspacing on tuber dry weight.

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Variety

120! I I

~ -. -El" I I I/ '8- _ -B- _ +:l- __ II/ i3---EJ

~

i90_!

I

!iI

6°-i

I

30. 0o Pentland Crowno Record

-120_ Physiological age

90.

"HOJ@ 30~HOJ

.g 08 -----------1.------ --.-- --r------- ---------r----.---

o Multio Apical

120

30

50

30 o 30cm

o 40cm

o ----TJune

-- ----------·1-----·----- -------r-July August Sept.

Figure 4.4.2.18 The effect of variety, physiological age andspacing on tuber number.

=<: SEDs[J

8

on

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Key: Q,Pentland Crown; 0, Record; open symbols, apical;closed symbols, multi.

60

C\J 50IS.. •HQ)

0~~ 40H •Q)

.gE-t

30~----~~------~~----~10 15 ~g

Stem number, m25

60

C\JI

S50

..HQ)

~ 40~HQ),.0~E-t

3035 40 45 50 55

LAD/TI30

Q

•800~.------~------r-----~125 150 175 200

LAD

Figure 4.4.2.19 The relationshipbetween total stem number (SN) (main& branch) ~nd total tuber number (TN).TN = 2.86(~8.45)+2.10(~0.42)SN%variance accounted for 77.4residual standard deviation = 5.12.

Figure 4.4.2.20 The relationshipbetween leaf area duration accumulatedfor a period of 30 days from the dateof tuber initiation (LAD/TI30) andtotal tuber number (TN).TN=10.44(~4.84)+1.32(~0.10)LAD/TI30for Record andTN=18.15(~7.23)+0.38(~0.15)LAD/TI30for Pentland Crown.% variance accounted for 98.8residual standard deviation = 1.14.

Figure 4.4.2.21 The relationshipbetween leaf area duration (LAD) andtotal tuber dry weight (TW).TW= 517.57(~123.58)+2.56(~0.75)LAD% variance accounted for 60.0residual standard deviation = 55.74 •

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7000

C\J •I 600E; •~0

"

!400MQ)

>0oj.)

~ TG = 205.72 (~ 74.66) + 15.82 (~ 4.42) LAD/AB• r-i 300 •Q)::t • % variance accounted for 62.8t: • residual standard deviation = 130.03.'lj

MQ).0 100e:!

o 5 10 15 20 25 30 35LAD/AB

Figure 4.4.2.23 The relationship between tuber dry weight over 57mm(TG) and leaf area duration contributed by the axillary branches(LAD/AB). For meanlng of symbols see figure 4.4.2.19.

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Variety ISEDs1250I

1000 I I

I .8-- --El150 »:

./I ...BI ./500.

:t: 0 Pentland Crown250 :I

0 Record

0

1250~Physiological age

1000

150........-:

.H

500C\JI 250So

0 Multi"~

0 Apical~ 08

_-El

1250 Spacing

1000 ___ -El

150 /'-Erc--

500 --:/'"250 ./0 30cm0 40cm0 --"-----_.-

TJune July August Sept.Figure 4.4.2.24 The effects of variety, physiological age and

spacing on total dry weight (TDW).

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Variety SEDs80

15

I_B--

/T/lY

'15

60L;.5

30 o Pentland Crowno Record

~-~-~-~~-~--~-r:------------.-~-

80'15

60~E--i

45Ct-i0

1j 300

1-1Q) '15,0;::!+'

* 0

Phys~ological age

o Multio Apical

80 Spacing

'15

6045l

30~15

o 30cmo 40cm

O·-l.---H=-----,r--------·T---------July August Sept.June

Figure 4.4.2.25 The effects of variety, physiological age andspacing on percentages of tubersout of total dry weight (TDW)(on dry weight basis).

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The effects of mix i t::; seed tubers of differentphysio-

Log i cc.L ~tge on: i.o. of dc~:ys from 9Lmting to tube r ini ti.at-

Lou ('rI) ('rIFP)j I.o , of d.iye from emer-gence to 'El ('l'IF~);

;:0. of duys from~)l:J.ntir:g to se ne scence ; (SEI':P) main stem

number (HSTL); Br-unch stem number (JJSffiJ) j number of axillary

brunche s (I.B).

TlFP rrr.: S·'c.:D ' 'cT" Dt"''l''l~~,,Jj, ... l"l0_~ .uu -2

m m m'I'r ea tmen t

Ap i c al, 53.6'(' n.3Lt 161.00 10.15 16.05 26.03

Multi 54.00 21.00 162.6'7 1'7.59 6.58 30.33

Cold 64.00 26.00 -:61.6'/ 1'7.28 5.11 1'?63

,\+t:: 25 53.67 20.67 158.00 1'7.65 1l~.38 17.65

A+I'l ;6 53.00 19.67 156.67 14.02 10.83 22.04

A+C 25 59.33 ~7 '2:7- 160.6'7 20.41 11.37 18.23C~) • ././

i;+C 36 58.17 21.03 161.67 12.42 9.60 25.32

H+C 25 5S.83 23.56 15'7.00 22.39 4.39 29.71

l·j+C 36 SE.oo 21.6'7 159.6'/ 18.33 3.44 23.27

SED 0.7'78 1.092 3.031 1.09 1.92 5.06

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In exueriment F3, 3 tubers of each treatment were dug up before

emergence to study the growth rate of snr-outs after planting. vii th

Table 4.4.3.2. The effects of physiological age arid sprouting treatments

on total sprout length after planting but before emergence (Exp.F3).

Total sprout Leng th, mm tuber -1

Date 21/4 30/4 21/4 30/4

Treatment Treatment

Apical 79.5 121 Fast 79.4 204

Multi 87.5 251 Slow 87.4 167

SED 13.68 58.1 13.68 58.1

availability of moisture and nutrients after planting, extension rate

of sprouts was increased in all treatments ('l'~ble1~.4.3.2 and Fig.4.4.1.2),

and so the initial difference in length of the largest sprout between

apical and multi of various experiments (Table 4.4.1.6 a,b) did not

affect the emergence. NumberE of main sprouts were greater in multi

and so this treatment had higher total extension rate of sprouts than

apical after planting (Table 4.4.3.2), while the higher extension rate

,of fast (Fig.4.4.1.3) was not maintained in the field (Table 4.4.3.2).As a result there was no difference in emergence due to sprouting treat-

ments (Fig.4.4.3.2). In case of multi the growth rate of sprout weight

was also higher (Fig.4.4.3.3) mainly due to a larger number of sprouts.

Due to increase in extension rate of sprouts, specific sprout weight

was decreased after planting in all treatments (Fig.4.4.3.3). Apicalhad higher specific sprout weight because sprouts were thicker than

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')3

those of multi. Mixing of different physiologically aged tubers or

different seed sizes (Bxp , F2, F4) did not affect the emergence. There

was no significant difference in stem numbers between different dates

of growth ar.alyses in all the experiments, and so they were averaged

for various dates of growth analyses until the middle of August for

after that they started to die. Like experiment F1, apical increased

branch stems, compared to multi in both experiments F2 and F3

(Table 4.4.3.1 and Fig.4.4.3.2), but the proportion of increase was

more in experiment F2 compared to F3 and total stem numbers were

increased in F2 and decreased in F3. Planting at closer spacings and

using bigger seed size incre~sed total stem number per unit area

(Table 4.4.3.1 and Fig. 4.4.3.1). 'Mixing of different type of seeds

and sprouting treatments did not affect the stem number or their type

(Table 4.4.3.1 and Fig.4.4.3.1). Numbers of axillary branches (AB)

were averaged for various dates of growth analyses after they stopped

increasing in number. Cold had lower number of AB compared to multi

(Table 4.4.3.1), this may be because emergence was more homogenous in

cold (Fig.4.4.3.1). Apical, in experiment F2 decreased the number of

AB (Table 4.4.3.1) while in F3 it increased (34.9,m-2) compared to

multi (25.4,m-2), this may be explained by the f3.ct that apical increased

the total stem number in F2 and decreased in F3, thus lateral buds in

F2 may have been ~haded more. On em overall basis over 90 per cent of

the AB present were contributed by the main stems which may be due to

two reasons: firstly, branch stems were lower in number (Table 4.4.3.1

and Fig.4.4.3.2) and secondly they were smaller in size (Fig.4.4.3.26),

and thus may have emerged later, so their lateral buds were shaded and

stayed dormant. In fact most of the effect appears to be due to the

sec.ond reason because numbers of AB con tributed by the branch stems

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were not in proportion to their total aumbers. For example in apical

(Exp. F2) the numbers of branch stems were more than the main still

main stems contributed aoout 85% of the totd.lAB present. This is also

supported by the fact that in case of mixing different physiologically

aged seed where cold VIas one of the compone nts , cold only contributed

23% of the AB present, because it emerged later, so its lateral buds

were shaded by the plants those emerged before it. As in 1979(Bxp , F1) a linear relationship was found bet\Veen leaf area duration

accumulated for a period of 30 days from the date of 50 per cent

emergence and the number of AB present (Fig.4.4.3.9). Data shown inthis Figure is from experiments F2 and F3 and every point is average

for 3 replicates.

4.4.3.2 Stolon growth and development

Cold and S25 increased stolon we igh t (Fig.4.4.3.4 and 4.4.3.7)

for a longer time and this may be due to delay in '1'1(Chapter 4.4.3.5).

Stolon numbers were :.I.lsohigher in these treatmen t.a. (Figs.4.4.3.5 and

4.4.3.7). Stolon gr-owth slowed down after T1 in :111 the three experi-

ments. l-lixingof differen t type of seeds and sprouting treatments did

not affect the stolon weight or their number (Figs.4.4.3.4; 4.4.3.5;

4.4.3.6; 4.4.3.7). As in 1979 (F1), a positive linear relationship

was found between LAl and stolon dry weight for all the 'three experi--1ments (Fig.4.4.3.8), until tuber dry weight of 19 plant was reached.

Data in Fig.4.4.3.8 is from all the three experiments and every point

is average for 3 replicates.

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4.4.3.3 Growth :md development of stem C:l.ndLeaf,

As in the previous year ma.i.nstems were slightly longer than the

branch stems, but overall plant height ',lasnot affected by any treat-

ment (Figs.4.4.3.10; 4,4,3,11; 4.4.3.12), except that early in the

season plants were smaller in cold and 825 treatments because they

emerged later.

Due to this their stem weight was also lower early in the season

(Figs.4.4.3.12 and 4.4.3.13). Increase in stem numbers per unit area,

either by planting at closer spacing or by using bigger seed size,

increased the stem weight per unit area, while other treatments did

not affect it (Figs.4.4.3.12; 4.4.3.13; 4.4.3.14; 4.4.3.15).

Increase in stem number [{Iso increased the LAI early in the season.

(Figs.4.4.3.16 and 4.4.3.19). Cold (Experiment F2) emerged later

(Fig.4.4.3.1) so had lower LAI early in the season compared to

Apical and Mul ti when considered from the date of planting, but

there was no difference when considered from the date of 50;C; emergence,

for example after 15 days of emergence multi had LAI of 0.7 compared

to 0.99 for cold and after 30 days of emergence multi and cold had LAlof 3.32 and 3.33 respectively. 825 (Experiment F4) also had lower LAlearly in the season (Fig.4.4.3.19), compared to other treatments in

-that experiment, which was affected in two ways: firstly it emerged

late and secondly it had lower total stem numbers (Fig.4.4.3.1).

In mixed treatments where" cold or 825 was one of the two components,

LAI early in the season was lower (Figs.4.4.3.16 and 4.4.3.19).

Competition between different components within plot is presented later

(Chapter 4.4.4.). Physiological age and sprouting treatments did not

differ in experiment F3 (Fig.4.4.3.18).

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After about 70 days from planting in all the 3 experiments LAI

was over 4.0 and effects due to different treatments disappeared

(Figs.4.4.3.16; 4.4.3.18; 4.4.3.19). In experiments, F2 and F3

senescence of the canopy was not significantly affected by any treat-

ment, thus there was no great difference between different treatments

in the decline of LAI. In eyperiment F4,S25 stayed green for 16 days

after other treatments had senesced wid thus had higher LAI later in

the season. For example after 149 days of planting it had LAI of 1.73

and B36 had only 0.25 (Fig.4.4.3.19). Although physiological age

(apical, multi) did not affect the LAI but the proportion contributed

by different types of stems was affected. For example in experiment F2,

main stems of apical, multi and cold contributed 62, 91 and 97}~of the

total LAI respectively. In experiment F3 these figures were 83 and 96

percent for apical and multi respectively. Difference between the

results of experiments may be explained by the different sources of

seed wh ich affected the type of stems Crable 4.4.3.1 and Fig.4.4.3.2).

Other treatments such as mixing of different types of seed (Exp. F2)

a~d sprouting (Exp. F3) did not affect the proportion of LAI contributed

by different types of stems. As in 1979, the size of AB was studied by

working out their LAI (LAI'AB) separately. LAI'AB was decreased by

cold and planting at closer spacing (Fig.4.4.3.17), these treatments

decreased the number of AB also. In Experiment F3, apical had higher

number of AB and thus had higher LAI'AB compared to multi (Fig.4.4.3.21)

while sprouting treatment showed no effect.

In gener-al,over 90 percent of the LAI 'AB in both experiments F2

and F3 was contributed by the AB present on the main stems. Number of

leaves present on the AB were proportional to LAI 'AB. Apart from leaves

coming from the AB, treatments those increased tota.l stem number also

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(1/

increased leaf numbers, for example, pl an t ing at closer spacing in

Experiments F2 and F4 and use of big seed size in Experiment F4

(Figs.4.4.3.22 and 4.4.3.19). Physiological age (apical, multi, cold)

did not affect the total leaf number except that cold emerged later and

thus had lower leaf numbers early in the season; but the important

result was that the leaves contributed by different types of stems wereaffected. In apical branch stems contributed a higher proportion of

leaves, wh i.Lein multi and cold most of the leaves were contributed by

their main stems (Fig.4.4.3.23).

Size of the leaves coming from the AB was not affected by any

treatment but their size \oJasmuch smaller than those coming straight

from the main stems. Thus average leaf size for main and branch stems

presented in Figure 4.4.3.24 was calculated w i thou t including the leaves

coming from the AB. In all the treatments leaves present on the main

stems were much bigger than those present on the branch stems. Leaves

"present on the br-anch stems of cold were srnal Leat i:l s ice, Differences

in leaf size resulted in differential values for leaf to stem ratios

for different types of stems (Fig.4.4.3.25) leaf to stem ratio decreased

as the season advanced due to increase in stem length. This ratio was

higher for main stems than for branch stems apparently because the

latter were smaller in size and so wer-emore affected by interstem

competi tion. Increase in stem number-s decreased the leaf to stem ratio

(Fig.4.4.3.2(7).

Specific Leaf area in Experiment F2 was only slightly affected by

interstem competition, increased by pLan t i.ngat closer spacings andwas not affected in Experiment F3. Thus leaf dry weight was prop-

ortional to their LAI (Figs.4.4.3.18 and 4.4.3.20). In Experiment F4

due to greater difference in stem number specific leaf area was increased

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with increase in stem numbers (Fig.4.4.3.19). As in Experiment F1,

the size of different type of stems was studied in Experiment F2 and

F3 and the results are presented as leaf area per stem (Fig.4.4.3. ~).

Main stems in general were much bigger than the br-anch stems, owing to

increased leaf size and the presence of higher numbers of AB. Apical

increased the size of branch stems compared to cold and multi

(Fig.4.4.3.26) and this difference was consistent throughout the season.

Mixing of different physiologically aged tubers or sprouting treatments

did not affect the size of specific types of stems.

Light interception (LVEarly in the season the effects of treatments on photosynthetically

active radiation (PAR) interception were similar to their effects on

LAI.

The relationship betweeriLAl and LI is presented later (Chapter

4.4.6). Cold and S25 intercepted less PAR early in the season because

they emerged later and had lower LAl. Increase in stem number due to

closer planting or the use of big seed increased PAR interception until·

about 65 days after planting. About 90 percent and most of the times

over 90 percent of the PAR waa intercepted by the canopy of all treat-

ments during the middle part of the growing season (65 to 120 days)

and differences dJ,leto treatments disappeured (Figs.4.4.3.27; 4.4.3.28;4.4.3.29).

4.4.3.5 Tuber growth and deveLopmerrt:

Tuber initiation (Tl) was calculated by interpolation from growth

analyses and additional sampling data, taken during early growth of the

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I)')

crop. Cold and S25 delayed TI for 4 to 5 d.iys when considered from

the date of 50% emergence. Thus apr-outi.ng tubers before planting

may have increased the tuberization stimulus. TI for mixed treatments,

in which cold or S25 was one of the two components, \oI:1.S calculated

separately for the different type of plants and then averaged to obtain

a figure for that treatment. Apical or multi sprouting treatments and

mixing of different type of seeds did not affect the date of TI.

In Experiment F2, cold increased tuber number over apical and

mul,ti and the effect \>I(1S more Lmmed iateLy after TI compared to final

harvesting (Fig.1+.4.3.30). This may be explained by the fact that LA!

at the time of TI in cold (2.69) was hi3her compared to apical or

multi (1.74), thus higher numbe r of tubers initiated but this difference

did not last very long due to better growing conditions (higher rain-

fall, Fig.3.3.1). As growth rate was very high, it did not take very

long to reach LAI of 4.0 when most of the incoming radiation was being

intercepted. In apical LAI of 4.0 "IDB reached after 14 days of TI

while in mul ti 3.11dcold it reached after 12 and 8 days of TI respect-

ively. Therefore assimilates available for the development of the

tubers after 14 days of TI may be similar in all the three treatments,

but cold had initiated more tubers and so the percentage survival of

tubers was more in apical and multi compared to cold. Similarly as

in Experiment F2 ~25 had higher LAI (2.8) at the time of T! compared

to B36 (2.40) thus S25 initiated higher number of tubers but percent-

age survival \'las.more in B36. Apical and multi did not differ sig-

nific~~tly, but multi had higher number~ of tubers and this differencewas consistent throughout the season (Figs.4.4.3.30 and 4.4.3.31).

Planting at closer spacings also increased tuber numbers which may be

attributed to the greater LAI at the time of TI (Figs.4.lt.3.30 & 4.4.3.32).

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1()!)

Grading of tubers showed that tubers which did not survive never attain-

ed a size of 25mm diameter and most of them did not attain the size of

10mm (Figs.4.4.3.30j 4.4.3.31; 4.4.3.32). Some wrinkled tubers and

f'ewin the process of shrivelling were found especially at the last

two harvests. As in 1979, a linear relationship between total stem

numbers (main + brwlch) and tuber numbers was found. Linear regress-

ion between them accounted 31% of the variance. S25 and cold appeared

different from other points, may be due to different physiological

s~atus of the seed.excluding only S25 from the regression, improved

the linear relationship (accounted 43.5% of the variance (Fig.4.4.3.}6) ).

Data in Figure 4.4.3.36 is from all the three experiments (F1, F2, F3)

and every point is ~~ average for 3 replicates. Since stem number do

not t3ke any account of stem size, thus regression equation between

tuber number and LAD, accumulated for a period of 30 days from the date

of TI accounted more variance (53.6%) and r-emoval,.of S25 from the

regression improved the liLear relationship (accounted 60%.variance)

(Fig.4.4.3.37). Data in Figure 4.4.3.37 is from all the 3 experiments

(F1, F2, F3) and every point is an average for replicates. Cold treat-

ment in Experiment F2 initiated tubers later than the apical and multi,

thus had lower tuber '{leight early in the season but when considered

from the date of TI there was no difference. For example after 12 days-2of TI, cold, multi and apical yielded 78.3, 88.7 al1d87.8g m respect-

ively. Similarly after 26 days of TI their yield was 352, 329.8 and-2 .326.6g m respectively. S25 in Experiment F4 also gave lower yields

early in the season, This effect was not only due to delay in TI but

this t.r-eatmen t also had a much lower LAl but then it senesced later.', than other treatments and so the final tuber yield vJUS not less.

Other treatments did not differ si.gni.f Lc.an t.Ly but planting at closer

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1 U1

spacing in Experiment F2 and F4 and api ce.L in Experiment F3 gave con";

sistently higher tuber weights during the growing season (Figs.4.4.3.31j

4.4.3.32; 4.4.3.33). As in Experiment F1 (1979), tuber weight was

related to LAD, wh ich accounted30)j of the variance (Fig.4.4.3.43).

Data in Figure 4.4.3.43 is from all the three experiments and every

point is average for 3 replicates. Tuber yield per unit of LAD was

4.3, which is lower than in 1979. This may be due to higher rainfall

during 1980 which stimulated more haulm growth. Tuber yield is usually

~elatedto LAD, calculated by assuming LAI over 3.0 as 3.0 (LAD'3)

(Bremner and Radley, 1966; Bremner illldTaha, 1966; Gunasena and Harris,

1968; Choudhury, 1980). 'I'hia relationship was examined here. Linear

relationship between LAD'3 and tuber we igh t accounted only 6 percent of

the variance and "'/asnot significant (tuber weight = 1038.57Ct 343.29) +

1.81(~ 1.28) LAD'3, RSD = 90.6). This may be due to the reason that

PAR interception was still increasing with increase in LAI over 3.0

(Chapter 4.4.6). Al though there was not much increase .in PAR inter-

ception with increase in LAI after 4.0, even than assuming the LAI over 4.0

did not give the significant linear relation between tuber weight (TW)

and LAD'4(T'v1 = 982.95(:' 310.16) + 1.64(:' 0.94) LAD'4, RSD = 87.89).

This may be explained by the fact that although there was no increase

in PAR interception with increase in LAI after 4.0 conversion efficiency

of the canopy was, improving (Chapter 4.4.6).

TNber size grades are of practical importance. Percentages of

various size grades for two dates of growth analyses in addition to

final harvesting are presented for all the three experiments (Table 4.4.3.3,

Figs.4.4.3.34 and 4.4.3.35). Similar results were obtained for other

.'. dates of gr-owth analyses not presented here. For simplification in

some cases tuber weight less than 35 or 38mm is not presented as their

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1U2

proportion was low (most of the time less than 2%) and there was no

difference between treatments. In Experiment F2 the cold treatment had

higher tuber numbers compared to apical and multi thus assimilates

available for the growth of individual tuber wer-e lower, and this

resulted in higher proportioLs of medium sized tubers. For examnle at

final harvesting cold had only 3.6 percent of tubers over 76mm wh iLe

this figure for apical and multi was 19.9 and 21.3 percent respectively.

The percentages of 38,57mm tubers \'Jere25.3, 7.0 Qnd 9.8 percent for

cold, apical and multi repectively. Per-centages of 5?...76mm tubers did

not differ significantly among these treatments.S25 in Experiment F4

also gave increased proportions of medium sized tubers (Fig.4.4.3.35).

Apical and multi of Experiment F2 did not differ but in Experiment F3

multi gave a higher proportion of medium sized tubers compared to

apical maybe because multi increased total stem number in Experiment F3.

Planting at closer spacing in Experiment F2 increased tuber numbers

and this resulted in a higher proportion of medium sized tubers

(Table 4.4.3.3). In Experiment F4, planting at closer spacing only

slightly increased the tuber number ill1ddid not affect the tuber size

grades. Tuber size over 76mm was related to leaf area dur3.tion con-

tributed by the AB, and linear regression between them accounted for.45% of

the variance (Fig.4.4.3.38). Data in Figure 4.4.3.38 is from experiments

F2 and F3 and every point is average for 3 replicates.

4.4.3.~ Total dry matter accumulation

Roots were not collected and those present on the stems and stolons

were removed and discarded; thus total dry wei.ght (Tm-J) figures which

are presented exclude roots. By the end of June leaves started to fall

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103

off and these were not collected from the ground. TmJ figures presented

in this chapter also exclude those leaves. Cold and S25 treatments

had lower 'I'm!because they emerged later. Planting at closer spacing

had consistently higher TD\'J throughout the season, wh i.Leother treat-

ments did not affect it (Figs.4.4.3.39; 4.4.3.41; 4.4.3.42). As in

1979, (Exp, F1) the ~ercentages of various components wer-e worked out

for these experiments. Cold and S25 recorded a lower percentage of

tubers out of TD'd (Figs.4.4.3.40 and 4.4.3.42) when considered from

tbe date of planting. But when considered from the date of T1 there

was no difference. For exumple after 12 days from T1 cold had 18.7

percent .of tubers, out of 'I'DHwhile this figure for apical and multi

was 20.8 and 20.9 respectively. Similarly after 26 days from T1 cold

had 45.8 percent of tubers while apical and multi had 47.2 and 43.9

percent respectively. Other treatments did not affect the percentage

distribution of 'I'DV!but apical of Experiment F3 did record a consist-

ently higher percentage of Dlv in tubers compar-ed to multi, but the

difference was not significant on any date (Fig.4.4.3.41).

4.4.4 Competition between two components within plot.

In experiment F2 there was mo intera.ction between apical and multi,

'so results are presented as sprouted and cold (average for apical and

multi). Similarly in Experiment F4 there Vias no interaction between

2 spacings used and so the results presented for that experiment are

averaged for 2 spacings. In all the three experiments statistical

analyses were car-ried out CtS split plot design taking two components,',

(sprouted and cold) within plot as sub plot and residual degrees of

freedom (RDF) is given on the figures itself. RDF for SED No. 5,6 of

Page 180: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

'I'abl.e 4.l~.3•.3. The ci I'e c Ls of mix.LIIG seed Lubers of different

physiologic:l!. itge or: pe r-ce n t,JGc of tubers in different

s i ze gr ..ide s out of to tal, tuber we igh t ,

D:J.;{s after 90 121 Final harvestingplanting

Size (mm) <35 35-45 <45 35-60 >60 38-57 57-76 >76'I'r-e a tmen t

A:pical (A) 2.5 15.0 82.5 11+.4 84.4 '1.0 72.9 19.9

Hulti (H) 1.lj. 18.0 80.6 211-.1 '15.3 9.8 68.7 21.3

Cold (C) 19.7 65.2 15.1 '73.6 24.7 25.3 70.4 3.6

A+E 25 4.0 14.6 81.4 42.9 56.1 17.6 72.4 9.5

1'.+1-1 36 2.6 20.1 7'1.3 24.0 75.2 7.6 75.6 16.5

A+C 25 11.3 16.7 72.0 53.7 44.5 23.6 68.7 7.0

A+C 36 10.3 18.1 '71.6 33.6 65.2 18.0 68.6 13.0

M+C 25 18.8 20.6 60.6 60.5 37.9 29.1 63.8 6.3-

N+C 36 10.1 21.1 6B.8 35.6 63.4 23.3 66.1 10.2

SED 3.28 5.94 6.22 '7.40 7.26 4.42 4.69 3.73

.'.

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40_

1III

32~I

i

Experiment F2o Apicalo Multib Cold

i

24!-"1

!I

15.Ji

I TI SEDs

HDate of 50% emergenceand its SED.

C\J --lI

s

I I SEDs...1-t 4o_, F4(J) Experiment

II0 Stem number

~ 0 B36~ 0 Averaged overi s: S25 I various growthS i

(J)

32J 0analyses.p B+S25Cl) I

i + B+S36 III

I

241 XlI

I161

lIII

8~

0 ,-L, r7 12 17 22 27 1 6May June

"

Figure 4.4.3.1 The effects of physiological age on emergence.

I

I

Page 182: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

C\JI

20,Ii

1SJiII

II12:

lI

8J

o Apicalo Multi

0" B·B 0

oc;6

I 0¢ po·oI 010

¥o

IT299 14 19

May24

Figure 4.4.3.2

fJ

o

o Slowo Fast

Stem numberaveraged overvarious growthanalyses.

20_.

rltil+'oE-tI SEDsoo

o

o1GJ

II

I

i12_;

9 14 19May

treatments on emergence .The effects of physiological age and sprouting

.-I 0 Apical

f-< 350_ .QJ ,0 Mu.lt ;.g+'

eo 280JS I

I

!+'ih'M 210 iQJ _,~

II

~ 0~ 70~E-t I planting

OJ_T~__ ~nthe f~ell

Jan. Feb. March April

Figure 4.4.3.3

QIIIII¢

o Apical

I3 0 GO, 0 Multi+ Slow*" Fast

3008.

~1 0 52~ T G-~

OJ1o ooL,___--.-- T_L~Jan. Feb. March April

The effects of physiological age and sproutingtreatments on specific sprout weight (SP8W) and the effectsof physiological age on sprout weight. Vertical bars areSEDs for last 3 dates and SE of mean for first two dates ofsmeasurements.

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\0 \0 \0(Y") (Y) (Y")

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Page 184: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

I=:0rl

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Stolon dry weight8.5 8.5

SEDs6.8 6.8

C\JIEleo..5.1 5.1

+>..c::bO'r-!OJ~>. 3.4 3.4~'CJ Apical>=1

0 0 Slow0 Multirl .0 0 Fast0 1.7 1.7+>U,)

Stolon number340 340

A I.I -,300 6 -, 300

-,C\JI 260 260El..~(J)

.02 220 -, 220>=1 'b0rlo·+> 0 Apical 0 SlowtQ 180 180

0 Multi 0 FastI

140 140 6June July Aug. June July Aug.

Figure 4.4.3.6 The effects of physiological age and sproutingtreatments on stolon dry weight and stolon number (Experiment F3) .

...

oJune July Aug.

O~-------'--------r-------June July Aug.

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.b- ___

- / ----- ---/ -~ - - ---- - ~._ - - - - - - - El/..... ------- ~-....;::::

// ------:_--::_

/'

II //

o ///x:' //

f{O-'-r---------·----··--T----··------ ----------1·-------June July August

Figure 4.4.3.7 The effects of mixing seed tubers of different Slzes onstolon dry weight and stolon numb~r (Experiment F4)

~/- 6 ./

/~

~o 00 L-.--_-----,----------r-- _.--...-1---1

000 045 090 1035 1080 2025

~ X10-1s1005tlO

" 084+'..Geo'.-1Q) 063~

t> 042<d

s::0rl 021_0+'UJ

000_

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Key:

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.b-........__./_./-- IX ........__~ SEDs/' A" ~G _ -/ /,' '~... ........_ -X

/ /' / "'-_ ____....._I Y/' -_;:.---~ - - __~

/ I f{ 'X'- - - ________I~ B

/ I

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II/

/

I I

/

o ,B36; /).,825; o ,B+825; +, B+836.

-:oy = - 0.686 (~0.274) + 3.555 (~0.25l)x% variance accounted for 85.4residual standard deviation = 0.741.

LAIFigure 4.4.3.8 The relationship between LAI and stolon dry weight.

Data is average for replicates and is from experiments: F2,qF3,AF4,0.

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I SEDs110

10

I0-

///

/~I#

I90

ElC)

50o Apicalo Multi

/

/1O......,_-------T----------- --- I

July

30

June August

Figure 4.4.3.10 The effects of physiological age on average plantheight (Experiment F3).

35'1Ii130 lI

C\J II IEl I

I.. 251p:j«: lt+-t

,

0

1-<. (J).0 '"~z

15120

,_)

y = 46.20 (~ 5.87) - 0.558 (~0.154)x% variance accounted for 50.2,residual

o standard deviation = 4.12

oo

o

-'--'-r ----.--r--'~~-~-T~---- ---,~--~---'I---~-I25 30 35 (,0 4'5 :SO ~):)

LAD/E30Figure 4.4.3.9 The relationship between number of axillary branches (AB)

and leaf area duration accumulated over a period of 30 days from thedate of 50% emergence (LAD/E30) . Data is from experiments: F2, 0 ; F3, J::,.

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ISEDsElCJ

105_ I

!0- - --------~:-?-::----~

.~ =r:>.cy .~-- .z>: _84 ," ----... .- ,/y-- -_/ -I .'.: G /'

63 I I //~- - ~-1 /-/- /

I /~ /421 f' /1 ",,;/ ."211 tt'/

o ~ _ -----1----------------- -- ---- --I ---------- ------------------,------

C\J

1Gb

250_

200_~I!

I

t.°lII

30_J

ISEDs

20~

-----------------T-------------------T-----July Aug.

Figure 4.4.3.12 .The effects of mixing seed tubers of different

size on average plant height, above ground (AGS) and under ground

(UGS) stem dry weight. Key: O,B36; b. ,825; O,B+S25; +,B+S36

(Experiment -F4) .

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en~Ji:l(f)

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Ul1-1

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Page 192: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

120 120

0 Apical o Slow~ 80 0 Multi 80 o Fasts~..CJl 40 40 .C,!)-<

20 20Apical 0 SlowMulti 0 Paat

~ 15 15El~..~ 10;::) 10

200Above ground stem(AGS)209

160160

o oJune July July Aug.JuneAug.

30Under ground stem(UGS)

30 .

25I SlIDe

25

5 5June July Aug. June July Aug.Figure 4.4.3.15 The effects of physiological age and sprouting

treatments on stem dry weight (Experiment F3).

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Page 194: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

rJlH

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LAl6 6

5 5

4 4 f

SEDs3 3

H

:!J2 2

0 Apical Slow0 Multi 0 Fast

o oJune July August June July August

160Leaf dry weight

160

SEDs

132

C\JI

1':1104eo"+>

~'(0 76~

t>'d~ 48cOQJN

o Apicalo Multi

7 DSlowo Fast

July

Figure 4~4.3.18 The effects of physiological age and sproutingtreatments on LAl and leaf dry weight (Experiment F3).

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Hj

I

II SEDs

July I August ..--T--·-------Sept.

550,C\J II IS f. f.o,

i" I

3301~._l~ ,~ 22°1Cri I

~ 1'10 iH -

I o ....o -,

T

, // /T, / /

/ /o / J!.)(/ / /

/s

....

....

IT SEDs

o_-!~~--------~---~~----~~~~-----June July August lSept~

:r4 I\\ fr - - - - _ -b- Ir--/~___ - -_.J:idI\ 7' c B - / - B - - - -:..._-_=-= - 8 - .::.-: --a

\ I : - £/ -~-

1/// //\/k

SEDs

1--June July August

.--r- -- ------Sept.

Figure 4.4.3.19 The effects of mixing seed tubers of differentsizes on LAI, leaf number . and specific leaf area (SPLA)(Experiment F4).Key: O,B36; b. ,S25; D,B+S25;+,B+S36.

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UJqI:Ll(j)

J I1

Ir-I+>

UJ::JbO::J~\0 \0 \0

(Y) (Y) (Y)

?f u u+ +

>.~ ~ ~30 0 <lt-:>

(j)

§t-:>

HI

ci-,c..) Cl C) CJ 0 0co I.II C\J Ul (J) CV),~ \~ ~

~+>UJ::JbO::J~If\ If\ ir-,

(\J (\J (\J~ u u+ + +< ~ ~0 0 <l

oDrloU

<l

,-.--------, ...- .. - ..- ......·T- ..···..- ..·-· -.,----, ....-- ..-- ..........------10000000co Ln (\J Ul (J) CV)~ ~ '\"

Cj...jcd(j)rl

s::0

UJ1-<(j)

.g+>oD(j)bOcd

>.rlrlcdo

'r-!eo0rl0'r-!UJ>.,.qp..

+>s::(j)1-<(j)

Cj...jCj...j'r-!oDCj...j0

eos::'r-!><'r-!SoD

~(l)eocd

~o

'r-!eo0rl0'r-!Ul

Ep..Cj...j0

Ul+> C\Jo Ii.(l)

Cj...j +>Cj...j s::(l) (j)s(l) 'r-!,.q 1-<E-t (l)

p..xIZ1

0(\J +>. ,.q(Y) bO

'r-!...:;t (j)

~...:;t

>.(l) 1-<1-< oD

61'r-!Ii.

Page 198: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

0 Apical0 Multi

1035_

ii

1008JI

!I

o 81~I __ BI yri - B-I I

o 54~p::) I~ /<,

IH

;j o 21J /

oooJ/

YJ-.-----'---,-- r -~.---.-.--- -T-···----June July Aug.

0 Slow0 Fast

'1035

1n 08.j I SEDs

o 8'1_:

o 54_J ,

o 21~I

n ool ----.---.-r------------l---·····---June July Aug.

Figure 4.4.3.21 The effects of physiological age and sproutingtreatments on leaf area index contributed by the axillary branches(LAI/AB) (Experiment F3).

o A+M25o A+C25b. M+C25

450_

C\JIS

385,IiI

32011I

I

~ 190JQ) , - IH I12St.

June..---r---··------

JulyI -

Figure 4.4.3.22

]85~

o A+M36o A+C36b. M+C36

IJ20~

!II

I1)55 !G 'l

!I

190~ I

125_1~ _, --r-June JulyAug.

I I

ISEDs

,---Aug.

The effects of mlxlng seed tubers of differentphysiologically aged tubers on leaf number (Experiment F2).

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54

(\J 210I S I

21G~!

Ii II152i

i!

'10SJ

5.4

Experiment F?

<,'N

-,-,

,R "/ \ -,\\\\

II SEDs

0._

//

.0./'

-:.e

./' --c-"'_./" -A---~- --7'1- ____0':--" ------- ..-_--- ·l-------------- .----T----·-·--~----

JulyJune August

Experiment F3

SEDsII

-_ -- -8-- - - - - -f=l- _ -_

---E)...... _---,-

Augusto c;:. "2"~-------Tn_u

June-'--1

July

Figure 4.4.3.23 The effects of physiological age (PA) on leafnumber contributed by the different types of stems, excludingthe leaves coming from the axillary branches. SED on the rightside is to be used for comparing means within same levels of PA.Residual degrees of freedom is 6 for experiment F2 and 8 forexperiment F3.Key: El,apical;O, multi; A,cold;--, main stem;---, branch stem.

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\ \

2025_Ii

1035

II10 8 O_~

Experiment F3

"Q)N-r-i "U)

1080_] SEDs

1D 35_iI!

-------------.---------------------a-August

Figure 4.4.3.24 The effects of physiological age (PA) on leaf size of thedifferent types of stems (leaves coming from the axillary brancheswere not included). For meaning of SEDs, symbols and residualdegrees of freedom, see figure 4.4.3.23.

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Experiment F2

2015

1062

o 0 0 ~---------------- --rJune

-TJuly August

Experiment F3.

~i,J 52

II1008

II II SEDe

June-----------------~----------_r

July August

Figure 4.4.3.25 The effects of physiological age on leaf to stemratio (dry weight basis) of different types of stems.For meaning of SEDs and residual degrees of freedom, seefigure 4.4.3.23.

Key: 0, apical; O,multi; 6., average for apical, multi and cold

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I21J

r-l 35_I

IEl(])+"u: I

C\J IEl 28~'d

I"cd(]) Ir-.cd ICt-i 211cd(])H

II

141I

Experiment F2

SEDeII

7 I I

___ -""- __ -6- _o ~--~ _...l..-;-if!F.:..__-------------r-------------- ----------------l-----~-- ---r

June July August

Experiment F3

II II B>s

.8-_/' -

/' - _ -L,/' r-"......._/' ---B' B- --- ___

/ -- -- -- --- --- --- --- ""El/ _B-- --s./ -_

o [t-3/ ---""0:---r~--------- ----------[-------------------- --_---I ------- ------------- __ ~

June July August

Figure 4.4.3.26 The effects of physiological age on stem Slze.For meanlng of SEDs, symbols and residual degrees of freedom,see figu~e ~.4.3.23.

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I1\\\ I\\\\-,-,-,

2 C(,D.) J....

IIi

I2 0 OO~

II

0

10501OJ I

~ 1000-1(j) I+' ,(f) I

050

June---'-',--'---..-...-.--·------ ..-·--·--1..--------"--'-'--'-'---'T---

July August Sept.

I

'2°i,

O-!-,r-------~~_.------_ ---r---- ---June July August Sept.

Figure 4.4.3.27 The effects of mixing seed tubers of different

SEDs

SEDs

sizes on leaf to stem ratio (dry weight basis) and photosyntheticallyactive radiation interception (PAR) (Experiment F4).Key: 0 ,B36; ~ ,S25; D,B+S25; +,B+S36.

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i!

8°1II

601

1I

IIII!40.~ ':

i 9, IDI i

s:: I 'I/ /o I' /'M I 0/;.> I _j~ 2O_-q~--------~-'---r'--'"'-_'---"'-"··-··-··-··-····-r··---··----··-·-·-----T----·--···-··HQ)

;.>s::'M

80~i

60JI

;

40J

I I

I ,k\,-Q I\ c::(>.0 'r] \

C'1qh\II/'

q5'p

SEDs

I.(

o ApicalIi . 0 M1Jlti

June July August Sept.

I'

!

f{({ o Slow

o Fast

ill/'.0/ /

I ~/20_~ _

June July August Sept.

Figure 4.4.3.29 The effects of physiological age and sproutingtreatments on photosynthetically active radiation (PAR)inte~ception' (Experiment F3).

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--I;UJ~['1(f)

H <1(.1 I J i:Q Ul

/) H~I (f)

!1J H s:::0

\/ / 0 Ul <0bO H I=:;j Q) ro1rrPQ t-i ~ .g H

I I \ -fJ Q)/

~r---t <1 r!) tp H <0Q)bO s:::-, '~ ro'\ :>, H

'fl cP~ 1-1 r-I :>, Q);j r-I .g\0 \0 \0

i II r;, r-I(Y) (Y) (Y) ro -fJ;2: 0 0.!;j.- $ H CJ+ + + 'r! r-I~ ~ ;2: -........ '-.- eo ro........---...'-. Q) 0 -fJ

0 0 <l .---....::.<;I-l § r-I 0B '<0 0 -fJ,------,------- ------,---- 1--------,------- r;, 'n

0 Cl) N CO <r 0 Ul H:>, 0r+ C"! 0 Cl) C"!~

ft-i\' \' \'

Cl)-fJ Hs::: roQ)

5:1U!=,+SaA.lBl{ TBU!=.if €JrJ HQ)~ft-i Ulft-i ~'n f:iI<0 (f)

QJ ft-i/ 0

I \ .:/ bOoI/ s:::

~ ~¢ bO 'n;j ~ I

\/\ ~ 'n '-'S

~~ ?:1~ <0I=: 0 C\J

1\ \ ro r-I r:r..:>,r-I Q) H -fJ

~ 0l\1 ;j bO Cl) s:::Lr\ Lr\ Lr\ r;, ro ::- Cl)C\J C\J C\J -< \ 0 s;2:' 0 0 .//

r-I 'n+ + + ::s- ----~ ro H H~ ~ ;2: CJ Cl) Cl)

-\:: : Cl) 'n~

~0 0 <l!!l .::-:::.--~ s::: bO

;j 0 f:iIr;, r-I s:::,- y--------r--------------r---------r---- 0-0 Cl) N CO -.....t 0 on H~Lt- C"! 0 Cl) C"! CIl Cl)

:>,~

0\' \' \'~ r-I

-fJH

ft-i <0 Q)0 s::: ::-

<l 00 til 0Ul

0 -fJ HCJ OJ

Y' Cl)

I ~ft-i

r4 Q) ft-iQ) s:::

-\ '-'Q) H

/ bO ..c: H OJ

Cy ~8 Q) .gQ

~-fJ

\ \ s::: H

~ ~0 0(Y) H ft-i

0 Q)

I / (Y)

~Q)

:>, H~ql r-I ..:;t -fJ tilr-I ;j 0ro 'n

\ 'i r;, ..:;t r-I~CJ -fJ <0 til CIl'n '3 r-Ils-_~ +J ~

~0 Q) 0 f:iI;2: 0 H -fJ (f),,- Cl)

So'0 0 <l § 'nr--------------!------------r-----------------,--------,----- r;, r:r..0 Cl) c\1 CO '.J 0Lt- (") Cl Cl) C"!\' \' \'

ill 'S.laqumu .la qn,r,G-

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1500

t\I11:11200~..~i900

~~ 600,Q.i:

90

75

60

45

30

15

300

o

Total tuber number (TTN)Tuber number over 10mm(TN010)

oJune July Aug. June July Aug.

Figure 4.4.3.31 The effects of physiological age and sproutingtreatments on total tuber number,tuber number over 10mm andtuber dry weight (Experiment F3).

..-?---

/ 0-- _ 0- __ 0 0/// 0

1/erI

J/1d

o Apicalo Multi

Aug.

o ApicalMulti

, 90

~..-I+>III

.-4 Q)cd tr:=•.-t 11fa4

SEDs (TTN)

75

45

15 o Slowo Fast

June July Aug.

Tuber dry weight SEDeo I ~1500o

1200

900

600

300 o Slowo Fast

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C\I 1700laI:VJ..1360+"

~'Q1 1020:.>..ff' 680fJ~ 340

200

160

120

80

40

200

160

120

80

40

o

/~....... I..... I SEDs/ .....__ , ,"'e-- - -_ - - - - ::...8-- ;;...:_-_A/..r--, __J"I _._ - :t::J

0-. , ca'''''' ...,4.~ __ • -- _..:: ---. __ .. __0--7 ~ ......_.--- -.,rr//

June July August sept.

I SEDs

SEnso•oJ:.

Figure 4.4.3.32 The effects of mixing seed tubers of differentsizes on tuber number over 10mm,total tuber number and tuberdry weight (Experiment F4)Key: 0 ,:836;J:. ,S25; 0 .BtS25; • ,B+8,36.

./I

June July August sept.

': ---~J:.

././

./

June July August sept.

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UJ>=lfLl(f)

I---l Cl it---t I

IL ~

0. Cf.leo 1-1;:::l OJ~ .g

~

+>"dOJeo~ '"3 ~

'0 '0 '0 ~ rl(Y") (Y") (Y") rl

=f u u '"+ + o~ ~ ::;<: OJ 'rl~ bD0 0 <I ;:::l 0~ r-i

0I

--______,- ----,--- ·rl0 ci 0 0 0 0 Cf.l

0 lD N 0) -0 E[1- CV! 0 lD CV! p..\' \' \'

+>:aU"F'+S<:JA.!Bl{ ~OJ

IBU!=.i1 0 0<1 1-1OJCriCri'rl"d

Cri0

bD bD;:::l ~~ 'rl~

'rlS"d~tll

~3 OJ

l!\ l!\ l!\ bDC\J C\J C\J ~ '"=f u u

+ + rl~ ~ ::;<: tllOJ o

0 0 <I § 'rlbD

~ 0I

--!-~'.-----,- ---,----- rl0 0 0 0 0 0 00 co C\J 0) '-J 'rl

Cf.l[1- CV! 0 co CV! E\' \' \' p.. C\J

Crif.%..

O@l 0 +>~Cf.l OJ+> So 'rlOJ 1-1Cri OJCri p..OJ ~~OJ

bD .cl;:::l 8 +'~ ~

'rlOJ

(Y") ~(Y")

~ ~3 (Y")

~rl ~ ...:::t

'" 'rl . 1-1() +> "d ...:::t OJ'rl rl rl '§Pi ~ 0 OJ~ U OJ S +>0 0 <I § bD

~ 'rlf.%..

0 0 0 0 0 00 co c\1 0) '-J •[1- CV! 0 co CV!\' \' \~

11& ''+l{:a::<:J.t1.A'!P .!<:JqnJ,G-

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<1

60 <1<1 <1

<1 <140

0 0 0 0 <1 <1

0 0 0 0 <1 <120 ..--•

0 0 0 0 ~ <1 ~ <l r-

• • 0 0 0 0 ~ <1 ~ ~ <l ic0

84 107 Final harvesting

+l'§,'r-! 80ID~

~"d

1-..ID'§ 60'+l

~

Fast80

Apical

40

20

o

Figure 4.h.3.34

Slow

Fast

Apical

o

o

Slow

Multi

o

Fast Slow

••

o 0

o 0

o 0

o 0

o

o 0

o 0

o 0

Multi

o 0

Apical Multi

•84 107

Days after planting

.---<l <1

<1 <l

<l <l

<l .<l

<l <l

Final harvesting

The effects of physiological age and sproutingtreatment on proportion of tubers in size grades (dry weightbasis). Vertical bars are the SEDs.[!] < 35mm 0 >45mm·· [§J >60mm ~ 57-76mmo 35-45mm @]35-60mm G 38-57mm lie I >76mrn

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+>ib 60.r!Q)

~t: 40'd

HQ)

,D

.B 20~

80

60

40

20

oB36

80

• •• •• •

91 days afterplanting (DAP)

825 B+825 B+836

o

125o 0 0 0 0 DAP

o 0 0 0 0

o

o

B36o 0 0 0 0

825 B+825 B+836o

60

40

20

oB36

Figure 4.4.3.35

<l

[!] <35mmo 35-45mm

D >45nnn

~ 35-60mm

o >60mm

~ 38-57mm

G 57-76mm

~ >76mm

825

<l

Finalharvesting

B+825

<J

<J

B+836

The effects of mixing seed tubers ofdifferent sizes on proportion of tubers in size grades(dry weight basis). Vertical bars are the 8EDs.

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average for replicates and is fromexperiments:F2, 0; F3, ~ ; F4,D . One pointencircled was not included in the

30T---'---'---'- __ 1I __ -r __~8 8 10 '11 12 13 14regression(seetext).

LAD/TI30 X 101

+:> 30 0 and leaf area duration contributed..c::bD'rl 25 by the axillary branches (LAD/AB)Q) 0~

-~ Data is for replicatest' 20 average'd and from experiments:NQ) '15 F2, 0; F3, ~.o 0~ 10

~A

53 4 5 6 '7 8 8 '10 1 '1

LAD/AB 'X101

y = 26.31 (+ 5.14) + 0.70 (+ 0.20) x% variance accounted for 43~5residual standard deviation = 5.08

50®55

°0

20 25 30 -2 35Total stem number, m

C\JIS

5055

50

y = 1.92 (+ 8.72) + 0.37 (+ 0.08) x% variance-accounted for 60residual standarddeviation = 4.27

®n

C\JI

51"~ y = -28.08 (~ 67.48) + 3.44 (+1.05)x

'te. X"" 0 '~i % v~riance accounted for 44.9 -N' J resi duaL standard~ 35 deviations=68.36o o

Figure 4.4.3.36 The relationshipbetween tuber number and total stemnumber (main + branch).Data isaverage for replicates and is fromexperiments:F2, 0; F3, A ; F4, D. One pointencircled was not included in theregression (see text).

40

Figure 4.4.3.37 The relationshipbetween tuber number and leaf areaduration accumulated for a periodof 30 days from the date of tuberinitiation (LAD/TI30). Data is

Figure 4.4.3.38 The relationshipbetween tuber dry weight over 76mm

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UJqr,lCl)

r-it----1 al

+>0+>

i .eo $:l

I ;:l 0I -<

~

[I)

~ID

'§+>

:>. ror-i ID;:l bDi-;) al

:>.r-i

\D \D \D ~(Y) (Y") (Y") ID V~ U U § OM+ + bD-< -< ~ i-;) 0

r-i0 0 <I 0OM

r ---I [I)

0 0 0 0 0 0 :>...c:::0 '0 CO N CO PlCO '0 0 r-- CV)

\' \' \' +>$:lID~IDr CHCH

IOM't1

bD

I ;:l CH-< 0

~

eo$:lOM><I OMa:>. ro3 $:l

i-;) al

IDeoalIf\ If\ If\

C\J C\J C\J r-i~ 0 U ID al+ + § o-< -< ~ OM

i-;) eo0 0 <I 0

r-i0I .--- OM0 0 0 0 0 0 [I)

0 '0 CO N 1O :>.CO '0 0 r-- CV) .g.'"' '"' \' C\J

CH r>:..0

+>[I) $:l+> IDv aID OM. CH ~

bD CH ID;:l ID

~-<ID rz:l

..c::: ._..,8 ..-...

~8:>. 0\

3 (Y)· +>i-;) (Y")

~·rl -:::t OMal OM · IDV +> ro -:::t ~0r-i rl rl

~ i 0~U ID ID

§ ~ ro0 0 <I

i-;) bDOMr>:..

0 0 0 0 0 0 00 0 I,.) () Cl ()CO I.n N CD 1O CV)\' \' ,-<

u6i '11G.10-

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UJ()~,jU)

H

ric0

m!--tQ)

I .g,I +''"dQ)eo

~ a:l

'3 ~I-:> rl

\D \D \D rl(Yl (Yl (Yl a:l~ () () tJ+ + + ·rl«: «: ~ Q) bO

§ 00 0 <I rl

I-:> 0·rli m

0 C\.i '0 CO 0 ~OJ ("1- In \' ..c:

P<+'s::Q)!--tQ)~'i-l·rl C\J'"d rx.'i-l +'

bO 0 s::~

Q)bO Ss:: .r-i.r-i !--t>< Q).r-i P<S ><

f;t:l'"d ..._..

~ §'3 Q) ~I-:> bO E-il{\ l{\ l{\ a:l

C\J C\J C\J

~() () fQ) rl +'+ + a:l ..c:«: «: ~ o bO

I s:: ·rl .r-i0 0 <I I~ bO Q)

0 ~rl_,--"---'---·'--1<>----. 0 t>0 N '0 co CO 0 .r-im '"dOJ ("1- \.() (") \' ~..c: rlP< a:l

+'~ 00 +'

r m ~+' 0oQ) +'

ibO'i-l ;J'i-l 0

~Q)

mQ) !--t..c: Q)E-i .g

+''i-l

0 0.:t

~ . Q)rl rl (Y') bOa:l .r-i ;J a:ltJ +' '"d I-:> .:t +'·rl '3 rl s::P< 0 .:t Q)«: ~ () tJ

!--t0 0 <I Q) Q)

Q)~ P<

§ bOI-:> .r-irx.

0 Lri 0 \.() 0 0OJ ("1- C..D '0 (") '\'

M.G.L JO q.no ' .Iaqnq. %

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TDW1520 1520

1220 1220

C\JI 920 920s

b.O

"~ 620 620E-i

0 Apical I 0 Slow0 Multi 'I 0 Fast

320 320

July Aug.

90% tuber, out of TDW

90

75 75

~ 60 60E-iIH0

+> 45 45;J0

Ulf..<OJ 30 30.g

Apical 0 Slow+-, 0

* Multi 015 0 15 Fast

June July Aug.JulyAug. June

Figure 4.4.3.41 The effects of physiological age and sproutingtreatments on total dry weight (TDW) and percentages of tubersout of TDW (Experiment F3).

SEDs

SEDs

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1850

1480C\JI s 1110

tU)

~6 140E-l

3'10

0

I I SEDs

-.-------...-------T-·----·-- ..---.- -r----·----------,---June July August Sept.

100

80~i:::)E-l 50~0

+-' 40::s0

.:rH 20Q)

.g+-'~ 0

:z:: SEDs

Sept.Figure 4.4.3.42 The effects of mixing seed tub-ersof different

sizes on total dry weight (TDW) and percentages of tubers,out of TDW (Experiment F4) ..Key: 0, B36; a , S25; 0 , B+S25; +, B+S36

X1011'10155

C\JI 150seo 155~+-' '150..r:::eo'MQ) -145~H 140Q)p::s+-' 135

30

o o

oo

y = 783.6(±262.5)+2.09(:!:0 ..14)X% variance accounted for 30.4residual standard deviation =7T~

·-A--r·····--·-·q-·· .._.--,--- ..-._.-r ···----r-·.--.-.---,

32 34 35 38 40 42UD X101

oo

Figure 4.4.3.43 The relationship between tuber dry weight and leaf area

duration (LAD)., Data is average for replicates and is fromexperiments: F2 (0); F 3 ( b. ); F4 ( 0 ).

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Figures 4.4.4.1 and 4.4.4.2 varied for different sampling dutes, but

there is not much difference in 'I' value (Table) for the range of RDF

found, so for simplification, average (for all dates) RDF along with

range is given. In Experiment F2 the mixture never yielded signific-

antly different from the exyected yield, i.e. the average of sprouted

and cold, when grown al.one (Figs.4.4.4.1 and 4.4.4.2). Replacement

diagrams for LAI and TD\v (Figs.LI-.4.4.1 and 4.4.4.2) show that competit-

ion between 2 components within mixed plot, started after about 60 days

from planting, ,8 sprouted in mixture guve higher values for LAI and

TDW compared to s9routed in mono and cold in mixture gave lower

values for LAI and TD'v'1 compared to cold in mono (Fic;s.4.4.4.1 and

4.4.4.2). Similar results wer-e obtained for tuber weight from 90 days

of planting onward (Fig.4.lt.4.2). In Exueriment F2 at 25cm spacings

differences between sprouted and cold were the same as found at 36cm

spacings (F.'igs.4.4.4.1,4.4.4.2 and 4.4.4.3). For examp.l,eout of total

LAI for mixture, after 63, 76, 90 :~~d 105 days of planting sprouted

contributed: 68; 64; 59; 60 percent at 36cm spacing and 67; 57; 69; 64

percent at 25cm spacing respectively. Similo.r results were for TmJ

and tuber weight. In Experiment F4 where difference between cold and

sprouted Wo.s further increased by using different seed size, sprouted

contributed about 90):; of the tobl yield of mixture (Fig.4.4.4.4).

During 1980, in both -the experiments sprouted hud an advantage over

cold early in the season but both componerlts senesced at the same time

i.e. cold did not show any advantage over sprouted later in the season.

But it was found that in Cl1se of Experiment F4, where cold was suppressed

more, the proportion of oversize (>76mm) tubers were ir.creased thus it

'may not be useful to have bigger differences. So in 1981 it was decided

to use· the S3lTleseed size with difference in sprouting only. Planting

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106

pattern W3.S slightly ch.mge d (Ch.ipte r 1+.3). There '..:.-J5 no interaction

between 2 pLanti.ngdates, and so the results presented are averaged

for 2 dates. Results for Experiment F5 sho\l that competition between

sprouted and cold was not affected by ch.mge in planting pattern as

results were quite similar to those obtained for Experiment F2 (Figs:4.4.4.5;

4.4.4.1; 4.4.4.2; 4.4.4.3). For ex~ple in Experiment F5 after 77, 98

and 119 days of planting (planting date tuken as aver:1ge of two plant-

ing dates), in mixture, sprouted contributed: 62; 63; 56 percent of

LAI, these figures are quite similar to those obtained for Experiment F2,

reported earlier.

4.4.5 CroD evaporation

In 1979, crop evaporat ion (ET) culculcted from the neutron probe

data \"as ah;ays lower than the potential evaporation (Penman, 1956)

(Fig.4.4.5.1). This may be rel3.ted to the lower LAI during that year

(Fig.4.4.2.8) and evapor-a t ion from soil surface may have been very

little as the soil was dry during most of the growing season (Fig.4.4.5.2).

For convenience the ratio of actual to potential evaporation (Fig.4.4.5.2)

is also presented which o.ppears to be increo.sing in favour of actual,

with a decrease in soil moisture deficit ili~da~ increase in LAI

(Figs.4.4.5.2 and 4;4.2.8). It shows that potatoes ar-e very sensitive

to drought. Another factor which may have contributed to the lower

values (calculated) of ET, is the capillary movement of ground water

\'Jhich\'Jasnot measured in the present experiments. In 1980 the field

had a slight slope, thus during he.ivy r-ai.nsdue to surface runoff and

drainage loss, ET could have been over-estimated (Fig.4.4.1.3), and

when the crop was aeneaci.ng, measured values wer-e lower than the

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3 2

--.-3.0

2.0

1.0

0105 121

H

j

0.5

153

3.0

2.01.0

o

Days after planting~igure 4.4.4.1 Replacement diagrams for LAI. The scale on left hand side

1S just for one-diagram.Key: 0, Cold;O, sprouted (average for apical and multi);

• , Mixture for whole plot; 1, SED to compare between sproutedand mixed (whole plot basis); 2, SED to compare between cold and mixed(whole plot basis); 3, SED to compare between mixed and mean of coldand sprouted (whole plot basis); 4, SED to compare between sproutedand cold within plot; 5, SED to compare between sprouted (half plotbasis) and its counter part in mixed; 6, SED to compare cold (halfplot basis) with its counter part in mixed. Residual degrees offreedom are: 16 for SEDs 1,2, 3; 8 for SED 4; 21 (17-24) for SEDs 5, 6.

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- --_. -- •1000- __

l?OO

750 900

500 600

C\i 250 300I

~~ 0 0"

121 139""''tb..,(l)~

t' 500'tj

~ 600(l) 200,Q 375;:J~

150 250 400100

125 20050

0 0 076 90 105

- -'-1000 --750 1110

500 740C\iI~ 250 370~"

""' 0 0fa 105 121..,&~'tj

"'itil

""'~ 40

Days after planting

600

400

200

o

Figure 4.4~4.2 'Replacement diagrams for total dry weight and tuber dryweight. For meaning of symbols, SEDs and residual degrees of freedom(RDF). see figure 4.4.4.1 except that RDF for SEDs 5 & 6 is 22(19-24) fortuber dry wt and 23(20-24) for total dry weight.

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Hj

I

SEDs

"..-_..f3'";a---

//

I/JZI

/

I

I SEDs850

C\JI '150seo"+J ~\ "·7 :~~\,.qeo

'rl(])~ r")~-

:>,,_)~ U

H'd

H 180(])

.g8

0 1--------

I

1-------- ---1---

1000ISEDs

I .oI --_B'"-C\J 800 B- -I ./s ./eo :t"./

" 600 01:: /eo /-'rl(]) 400 :1:/~

'~..0-:'d 200 =r,/

rl gcd+J /'0 /'8 0 _-_"---

June July August Sept.

Figure 4.4.4.325cm spacing.Key: 0 ,sprouted (mean of apical and multi); 0, unsprouted

Growth of 2 components within mixed plot at

(cold)(Experiment F2).

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1r;nn/0... ... _.o

..../ ...... 420

N / 'DNI IS 1200 / I seo /

bD

+> ;:f +> 280.t:: .t::

bDeo 800 / 'rl'rlQ) p/ Q)

~ ~

t: / :>. 140H'd 400 / 'd

M ;1 MtU 0 tU+> +>0 08 80 0

1300 ID- ---0 350 )J/ I -:

/ INN p/I ISS /bD /

bD /800

+> / +> 210.t:: .t::bD eo'rl / ·rlQ) OJ~ 0 ~

/ :>.:>. 400 / IH H'd rf 'd

H / H 70Q) OJ

,..0 / '§;:J IT8 80 0

Figure 4.4.4.4 Growth of2 components withinmixed plot (Experiment F4). Residualdegrees of freedom (RDF)is 4 except for 4th sampling for which it.is 2.'Key: 0, Big; 0" small. Vertical barsare the BEDs.

2.0

,I'0

1.50

1.0

0.5

oJuly Aug.

Figure 4.4.4.5 Growth of twocomponents within mixed plot(Experiment F5). RDF is 4.Key: 0 , apical; 0 , co.Ld .Vertical tars are the SEDs .

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1(),/

Pote nt i al , which is r eLute d to dec.line in 11'.1. For- further calculations,

Potential evapor a t i.on data for 1980 and me.rsured for 1979, are used.

Evaporation for var i.ous t re a tment s of Experiment F1 (19'79) is

presented in Fig.4.4.5.1. Evupor-c.t i on Wi,S only slightly affected by

varieties. Apical t.r-eotrnent had J. slightly higher evaporation rate early

in the season but Late r on multi had hi gher compar-ed to api cal , For

example, total, evaporat i.on (HJ.i) from 4th June to 30th July and 31st July

to 7th October W:IS '78.5 and 93.2 for ,J.piccc: and 71.0 .md 101.2 for

mul ti respectively. Sp.ici.ng did not ..if'f'e c t the evupor a t ion (Fig.4.4.5.1)

except that during the L...ter par t of the growing season evapor-at i on was

slightly higher in s40. During severe drought roots of the variety

Record penetrated sligh tly deeper than those of PentLand Crown (Fig.4.4.5.3),

other treatments had no effect on rootir:g depth (data not presented).

In 1980 before the onset of he avy ra ina the soil IvJS quite dry early in

the season ar;.d roots by 11th June were extr:.tctinG·water fror:J 70-80cm

depth and there W:J.S no difference amongst the various treatments. After

that rocit penetration could not be followed as most of the time the field

was near field capacity. At the end of the season (beginning of Sept.)

it was found th:.tt roots did not go deeper th.,n SOcm.

The re~a t i orish i.p be tween crop evupora t ion and crop gr owth rate is

presented in Figure 4.4.5.4. Leaf senescence w i thin the canopy ','las

evident in 1980 .rt .n e.u-Lier date Lh.in Ln 1979, vrh i ch may have been due

to differences in humi.di ty .rnd L1\l between the year s , Consequently, the

t.otaL dry weights for 1980 were udjus ted to ...ccount for the early leaf

loss, w i th estimates of led weight loss being related to the dry weight

of senesced Leave s and number of missing Leave s , CrOiJ growth rate data

presented in Figure 4./+.5.4 is'ldjusted (roots not LncLuded ) and refer

to harvests up to 15th August in 1980 and 20th August for Record and

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108

10th September for Pentland Crown in 19?9, since it vJ;J.S r::ot possible to

account for weight losses due to stem rotting after these dates. The

five points encircled on the Figure 4.4.5.4 were not included in the

regression, four were for the ye~r 1980 when ground w~s not covered

and thus Potential evaporation \HS over=e s t iraate d and one point for 1979,

Pen tLand Crown, 27th July to 8th August, a.s a lot of the leaves had

senesced during that period 'J.nd no adjustment was made. Linear re-

gression for the remaining p~ints accounted for 79.5 nercent of the

var-Lance in crop growth r:J.te ,

4.4.6 Light interception ::l..ndpot:lto growth

4.4.6.1 Led are:" index :'\Ddphotosythetic:llly active r<'_diation (PAR)

intercention.

The relu.tionship between leaf :.trea index (LAI) cmd the proportion

of P1Jt intercepted by the crop canopy is shown in Figure 4.4.6.1. Up

to LAI values around two the relationship appeared to be almost linear,

wh.iLe above four, PARinterception W"1.S relatively constant. The charact-

eristics of light transmission in crop ccnop i.es have been related to

Beerl's Law by the equation IL _ 10 e -kL wher-e IL = the irradiance on

e, hor-i zon tal, plane below a leaf are a index of L, and k ,. a light ext-

inction coefficient, assuming n homogenous canopy, which depends on the

transmission chur-ac ter-Ls t i cs of single leaves and their geometrical

arrangement (Nonte i th , 1965). The r-eLat.Lor.ahip generated by this

equation did not fit the measured d~ta over the range of LAl found in

these experiments (Fig.4.4.6.1). From the analysis it ilppeared that the

light extinction coefficient v.il.ue , k , W'clS not constant as LAl increased.

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Varietyb. Potentialo Pentland Crowno Record

) ."C.~ cf._!_

,.'" .-)t::",,~ D ,~ •. )

30'15~I

~3025

']j

~ 20'15"s:: 20250

'rl~roH 10150p.,

~QJ 1025p.,0H

o 75u

Physiological age

o Multio Apical

~/' .B- -B' \

\ I1':1

-----··--J--·------···--·--r----·--- ·-------·-T--------

30 j5~ Spacing

30252075202510151025

o '15_.June

Figure 4.4.5.1

o 30cmo 40cm

I

,R

/ \\\--fj

July .I ·····------·--T-·-···----·-August September

The effects of~riety, physiological age andspaclng on crop evaporation (Experiment F1).

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~ 56

090,II

Io 19,

I

o68JI

10

28,

1:]I---.-------.I-----.------.T----.-- ...--.----.-----rJune July August Sept.

I '03 5---·------·-·-·--·-r---·---·-·---.-------r---- ....-.-.---.-T

June July August Sept.

Figure 4.4.5.2 Cumulative soil moisutre deficit and ratio ofmeasured (neutron probe) to potential evaporation (ExperimentFl, average for 2 varieties).

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0

0 Pentland Crown20 0 Record

Elo....c: 40+'p..(lJ

'd

eo.:.: 60.r!

+'0 \0p:;

~

80 \\b-e

100June July August Sept.

6. Potential evaporation3.75i Crop evaporation0

3.1~I

I I \~ 2.4 I ~'d

~ J. \\

I \../.:.: \0 1.8 I'r! \+'

al 6.J...t0c,al::-~ 1.1

June July August Sept.

Figure 4.4.5.3 The effects of variety on rooting depth(Experiment F1) and crop and potential evaporation during,1980 (~xperiment F2).

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~I>. 20!O'0

C\JI-etJI..t3u 10

o

Y = 10.98 (!1.27) x - 2.56 (!2.48)

% variance accounted for 79.5residual standard deviation = 2.78

1.0 2.0-1ET, mm day

1.5 2.5

Figure 4.4.5.4. The relationship between crop growth rate andcrop evaporation (ET). Key: 0, exp , F2; ~ ,exp. F3;o ,exp. F4; • and • ,exp. F1 (Pentland Crown andRecord respectively). The points encircled were not includedin the regression (see text).

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109

In these experiments, Lncr-e ased LAI was ussociated wi th crop gr-owth

as a function of time and therefore it may be expected that changes

in leaf angle, size, distribution and transmission characteristics

with leaf age 'tlouldoccur. A significant (p = 0.001) quadratic re-

lationship was found between k and LAI and could be represented by the

equation; k = 0.335 + 0.1511 - 0.013L2 (Fig.4.4.6.2) (where L = LAI).

\oIhenthese calculated values for k were substituted into the Beer's

Law equation the resulting fit to the data accounted for 9~;6 of the

variance compared with 88% when a constant k(= 0.72) was used

(FiJ.g.4.4.6.1). Thus the du t a indicated that k VIas relatively lower at

lower leaf area indices, earlier in the season. Differences in leaf

angle which may occur during the season would obviously affect the

light interception characteristics of the canopy, but such differences

wer-e probably confounded wi th other changes for example leaf distribution,

and individual effects could not be determined, from these experiments.

The PAR interception da1.;u(Fig.4.4.6.1) was obtained from all the

treatments of Experiments: F1, F2, F3, F4 and although LAI varied, the

relationship betweeriLAI and PAR interception did not vary systemat-

ically with any of the agronomic treatments.

4.4.6.2 Dry matter nroduction

Daily incoming PAR data was obtained from a site within one km. of

the experiments (courtesy of H. Bat.eman, Environmental Physics Section).

On a few occasions when instruments failed, total solar radiation CST)

values were obtained from u.neu.rby Heteorologicill Station and PAR was

calculated as PAR = 0.53 (ST), from the relationship shoi...n in Figure

4.4.6.3.

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A significant Li.ne ar r-eLat ionsh i.p was found between cumulative

total dry weight (TmJ) (1'D'.1,used in the chapter is adjusted see

Chapter 4.4.5) and cumulative PAR intercepted in both years, and the

1980 data are shown in Figure 4.4.6.4. 'i/hen forced through the

origin, total dry weight ('rm-J) = 2.49 (PAR) and 3.42 (PAR) for 1979

and 1980, respectively, .indi.ca t ing t.hat conversion of light to dry

matter was more efficient in 1980. Al though such iJ.E effect may be

explained in terms of wate r stress in 1979 (Chapter 4.4.5), relation-

~hips based on cumulative data are not entirely satisfactory since

other variables, and time, m.iy be confounded with the apparerrt effect.

The photosynthetic conversion efficiency (g dry wt MJ-1 PAR

intercepted) for the canpoy W:lS cul.cul.at.ed for each gr-owth analysis

period. In gener a.l , the expe r imen'to.L treatments had no significant

effect on photosynthetic efficiency and therefore the general temporal

patterns for var-Le t ies in 1979, and individual experiments in 198o,

are shown in Figure 4.4.6.5. In general, photosynthetic efficiency

was higher in 1980 compared with 1979, probably as .':1. consequence of

differences in water stress. The variety Record had a higher con-

version efficiency than Pentland Crown, during early August, and it

may be that Record was less affected, since neutron probe data indi-

cated that Record had a slightly deeper root system. Later in August

the photosynthetic efficiency of Record vIas Lowe r than that of Pentland

Crown due to earlier C:_U10PY senescence.

Crop gr-owth rate (CGR) ~lS a function of PARinterception is shown

in Figure 4.4.6.6. A Li.near- relationship was evident for 1980, while

CGR Wi.l.S severely restricted under the drought conditions of 1979.

Calculation of intercepted PAR, assuming constant le = 0.7, from LAI

values resulted ina similar relationship between CGR u.nd PAR.

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111

However the r-eLut i.onsh io vii th measur-ed PAn accounted for 91% of the

var i ance in CGR..rh iLe th.i t with calculated ?1\R intercention accounted

for 84%. The difference was due to the f',ct that k WCJ.6 not constant

throughout the season dS discussed previously.

Al though CGRand to ta.l dry m.it te r production are .impor-t an t the

factor of major COnCer!l in pot~toes is tuber yield. The relationship

between tuber yield and interceuted PAR is shown in Figure 4.4.6.7.

-1Although photosynthetic conversion efficiency (g tuber dry wt HJ PAR

i_ntercepted over the whole season) for tuber weight was slightly

different for two years: tuber dry weight = 1.97 (PAR) and 2.37 (PAR)

for 197q and 1980 respectively, but still Q significant linear relation-

ship between tuber dry weight and to t al, PAR intercepted over the whoLe

season existed CFig.4.4.6.7). Between two varieties in 1979, Record

vias 13.4% more efficient than PentLand Crown in converting light to

tubers.

4.4.7 :Sxperiment F5

Soil temperature, at 10cm depth lnd screen, max. and min. air

temperature and rainfall from April to October 1981 are given in

Figures 4.4.7.1 and 4.4.7.2.

4.4.7.1 Emergence ,md stem number

The cold treatment took 45 days to reach 5CJ;G emergence compared

to 31.7 for upi cal., :, dd ff'e r-ence bigger th:m the one f'ound for Exp, F2

C1980). This muy be exp'lairied by the difference of temperature at the

time of p.lan t i ng of two t r-ea tmen t.s in this exper-Lment , Similarly

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Figure 4.4.6.1 The relationship between LAI(L) and interceptedPAR in 1979 and 1980. Each data point is the mean ofreplicates for a particular treatment. Excludes data fromlate in the 1980 season when LAI was declining and stemswere intercepting a higher proportion of radiation (totalnumber of points 278).

(a ) p. . 1 -kL h 0 2 ( )a AR lnterceptlon = -e ,were k = .7 ----residual standard deviation (RSD) = 7.90% variance accounted for 88.26.

(b)-lkLPAR interception = 1-e (----) 2

where \ = 0.335 (~0.038) + 0.151 (~0.030) L-0.013(~0.o05)L

RSD = 6.49% variance accounted for 92.1

Key: 0, Experiment F2; D. , Experiment F3; 0 , ExperimentF4; <:> , Record /Experiment F1); \l , Pentland Crown(Experiment F1).

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o

o

<loo

o

o

oI>I>I>o

. I>-'80. 0c»- 1>1> I>. Rf 0I> ~ 1>0 I>OD'l-""OO~ "

I> 0~~t>\o ' 0 0

I> ,1>0I>oI>

®:>O0:I>

o o

I> 0

"o

oo

o

1O

o

If)

CV) H

j

(\)

r---r----.---,----T----r----.-----T--_.-- __ ~--_,__--_r 000 ~ W If) ~ CV) ~ 0

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PAR = 0.53 (~ 0.001) ST% variance accounted for 99.9residual standard deviation = 0.20

+-----------1------- -

5--- r------ ------------ -1 - 1 -------

10 '15 20-2 -1ST, MJ m day

The relationship between photosynthetically active radiation

_ .. -.. _---o

30

Figure 4.4.6.3(PAR) and total incoming radiation (ST) on a horizontal plane. Data from144 days between May-October, 1979.

Co

X10-110_

9--,:

8:..,'1j

i5~sJ

I

k 4J 0i

3J

')\.-' -,.\"0

,,.r-,

oo

2k=O.335(~0.038)+0.151(~0.030)L-0.013(~0.005)L% Variance accounted for 42.2residual standard deviation = 0.14

+r2

rJ

r------------- -----,-------

4 5i

5L

Figure 4.4.6.2 The relationship between k and leaf area index (L).Data from experiments: Fl; F2; F3; F4. Excludes data from late lnthe season when LAI was declining and stems were intercepting ahigher proportion of radiation (total no. of points 195).

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I\-t0

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30 CGR = -2.97 (~1. 56) + 3.85 ± 0.28) PAR% variance accounted for 91.3residual standard deviation = 2.48 0

D

r- 2I

~c

-0C\J 15I v

AS o oeo

0.. Vp::0 c0

5

00 2 3 4 5 6 7 8

PAR, MJ-2 day -1m

Figure 4.4.6.6. Crop growth rate (CGR) as a function of PAR interceptionfor three experiments in 1980 (each data point is a mean for totalnumber of plots in that experiment) and 2 varieties ln 1979 (each datapoint is a mean for 12 plots). Regression line fitted through 1980 dataonly (see text).

1700TW = 428.71 (~134.74) + 2.98 (~0.23)PAR% variance accounted for 87.9

OAresidual standard deviation = 103.03.

C\JIS

eoAD

+'..c:bO'MQ)

~»H.-0

VHQ) V.£l~E-i

80650 70

Figure 4.4.6.7PAR, intercepted MJ m -2

Total tuber dry weight produced during 1979 and 1980". as a function of PAR interception. Each data point is a mean for

replicates .

. Note: For meaning of symbols see figure 4.4.6.1.

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ear Ly planting :.llso took more di_lys to r e ach 50% emergence compared

to late due to differe~ce in temper~ture. (Table 4.4.7.1.). Unlike

Experiment F2, cold took more days (8.7) from appearance of the first

stem to reach 5(J)b emer-gence compar-ed to ap i.caL (6.l,), this may be

due to the r eaaon the, tJ.pic21 in Expe r-i.ment F2 .incr-eaae d tota.l stem

number. Stem numbers were averaged for .dL dates of growth analyses,

as there was no difference be twee n different da te s and they were not

uffected by :my tr-ea tmerit (T,Jble 4.4.?1.).

4.4.7.2 Growth and develoument of stem and le.::.f.

Cold emerged later ur.d thus hnd lower LJI,I ear Ly in the season

(Fig.4.4.7.3) but when considered from the date of 505b emergence cold

had slightly higher LAl compar-ed to .ipi.cnl , For exampLe after

14, 21 and 28 days of 50}Semer gence , cold had LAl of: 0.l~2; 1.14 ;

1.65 and apical: 0.40; 0.86; 1.34 respectively. Late pLant i.ng had

lower LAl early in the season but when cons ider ed from the date of

50;:; emer-gence there was no difference. For example after 12, 19 and

26 days of 50}S emergence Lat e had LAI of: 0.53; 0.97; 1.51 and early:

0.43; 0.87; 1.54 respectively.

As found for Experiment F2, mixture gilve s ign i f i.cunt Ly lower

LAl early in the vseason cornoar-ed to .rp i cu'l , Specific leaf area

was not .rf'f'e c t.ed , thus leaf dry weight vl:AS pr-cpor t iouul. to LAL

Effects on stem weight ..us s irni.Lur to LAl. Leuf to stem ratio de-

creased during the gr-owing se ason :lS wus found in the previous two

years and w:J.snot af'f'e c t.ed by any of o'~hetreatments (data for

leaf and stem weight is not presented).

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113

4.4.7.3 Tuber growth :md developmen t ,

Tuber Lni t ia t ion (TI) W.J.S cal cu'Lrted by interpolation aa in

previous years. Unlike 1980, TI was not delayed by cold (Table 4.4.7.1.)

when considered f'r-ornthe date of 50;\ emergence, but when considered from

the appe ar ance of first stet .., it vias 2.7 days La te r in cold compared

to apical; still the difference was slightly less then the Experiment

F2 (4.3 days). Tuber weight was lower in cold and early planting

(Fig.4.4.7.5) but when considered from the date of TI there was no

difference. For example af te r 5, 15, 36 and 57 days of T1 cold yielded:

6 4 41 6 -2 . 1 2 102 8 3 8 6 -23.7; 92.; +2; 51gm and ap i ca.i : 1 .1; .; 7; OOgm ,

respectively. Similarly iT: case of date of planting (data used only

from mono plots as da te of TI for mi.x tur e vlilS not worked out), after

6, 21, 37 and 58 days of TI, late plonting yielded: 33.6; 110.5; 416.5;

580.5g m-2 and early planting: 28.'7; 122.7; 422.8; 611.2C m-2 respect-

ively. As found for Experiment F2 (1980) tuber weicht in m::'xture was

significantly lower than apical ear-Ly in the season.

As far as tuber size grades ,ITe concerned cold did increase the

percentage of medi.urn size tubers (Fig.l+.1~.'7.6) but the difference was

not significant. Late planting had a higher percentage of medium sized

tubers, but maybe because it was t.r-a.i Li.ng about 4 days behind (difference

in time to rea.ch501{' emergence), early pl:mting.

As found in 1980 (Exp, F2) cold had slightly higher LA1 (1.30) at

the time of T1 then apical (0.98) and Lni t.ia ted ~l higher number of

tubers (Fig.4.4. 7.4). Planting date did not affect the tuber number-,

More tubers were found in r:Jixture compar-ed to api ca'l , maybe due to

presence of cold in the r:Jixture.

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4.4.7.4 Tot3.1 dry matter ac cummul.et iou,

During this year roots present all the stolons iJnd stems were not

removed. Leave s started to falloff, by the middle of July and were

not collected. Like LAI and tuber we ight , total dry weight (TD\-l)

was also Lowe r in cold and late plan ting (Fig. 4. 4. 7.7) due to diff-

erence in time of emergence (Table 4.4.7.1.). Percentage of tubers

out of TDHwere lower in cold and Ls te p.l an t Lng (Fig.4.4.7.7) but

~hen considered from the date of TI there was no difference. For

example, after: 5; 15; 36; 5'7 days of TI cold had : 16.2; 35.3; 60.5;

68.6 percent and apical: 9.25; 32.94; 57.38; 68.3 percent tubers out

of Tm-l, respectively. Similarly in the case of date of planting after:

6; 21; 37; 58 days of TI, late planting had: 16.4; 34.5; 59.3; 67.0

percent and early planting had: 14.7; 3'7.0; 60.4; 70.6 percent tubers

out of TD\ol,rcspecti ve Ly (dab used from the mono plots only).

4.4.8 Exneriment F6

This experiment was specially designed to study the effect of

physiological age on emergence. 'I'her-e was no difference in time to

reach 50;; emergence between BODand 920D suggesting tha t once. the

sprouts had become visible (ubcu t 2 to 3mm) af te r- that Longe s t sprout

or phys i.ol ogi.cal. age has not much to do with emergence (Tables 4.4.8.1.

and 4.4.1.6.). Time between appearance of first stem and reaching

the 50;; emergence var ied from 2 to 6 days and VIas 4.0 for 4D and 5.5

for 920D.

All the plants Vlere harvested after 64 days of planting. Although

LAI vIas higher in treatment 920D it did not differ significantly from

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the treatments which emerged with it (rI\~bles 4.4.8.1; 4.4.8.2.).

Similer results were obt.a.i.ned for stem wei ght , Leaf weight and total

dry weight (Tm'!) (T,',ble 4.4.8.2.). Tuber weight , tuber number and

percentage of tubers out of TD'.-J, were hi ghe r in tre a trnent 920D

(T:J.ble 4.4.8.2.) pr-obub.l.y becauae it initiated tubers earlier. But

all other treatments of those which emerged at the sarne time did not

differ significantly (T;3.ble 4.4.8.2.).

Table 4.4.7.1. The effects of physiological age and time of planting

on: stem No, , emer-gence and tuber initiation (TI).

Days be tweenTotal Days taken appearancestem to reach Days taken of 1st stem

nu~~er 50}; to TI from to reachingm er.Jergence planting 50,;6emergence

Treat-ment.

Cold 14.9 45.1'1 68.17 8.69

Apical 13.6 31.6'7 54.33 6.35

A + C 13.4 40.0

, SED 0.68 1.322 1.139 1.139

Early 13.3 1+0.?8 63.33 8.19

Late 14.'1 3?11 59.17 6.85

SED 0.55 1.088 1.344 1.139

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100

60

~n

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""""PI,,..,oQ)s-.a,

80

-

-

-

-

-

-

,-

20

o April May June July Aug. Sept. Oct.*

Figure 4.4.7.1 Monthly rainfall from April to October 1981.

*only for first 25 d~ys for October.

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XBUI Has -_ -~-- --3>

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Page 244: Khurana, Subhash Chander (1981) Growth, development and ...eprints.nottingham.ac.uk/13729/1/256347.pdf · ACKNOWLEDGEMENTS I am greatly indebted to Professor J. D. Ivins and Dr. J.

015

I ISEDs3 n '15

- --El

-8

I./

10I

o Apicalt::.. Coldo A + C

/II \':o //

/ ;or

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015

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'////

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----0

o Earlyo Late

o 0 O,...l-.,- ~, ,_ " ~-, _'I'

June July AugustFigure 4.4.7.3 The effects of physiological age and date of

planting on leaf area index (LAI).

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100 I I I SEDs100

r ISEDs80 80 I Ja._

I ' 0----0 ....,.., -.......,

C\J 60 m..... 7I !Pi .-, <, ,- ..D 60a I I' ,,-- /n

I I "Et'I /H 1 ¢OJ 40

m /s::: o Apical 0 EarlyH /OJ 20 A Cold o Late.g 20 I8 0 A + C 13-~

0 0June July Aug. June July ug.

Figure 4.4.7.4 The effects of physiological age and date of planting ontuber number.

650ISEDs ISEDsfLI 550//

1//

380I I

,

I ~ 260II 0 Apical 0 Early!Zf A Cold " 0 Late

/ I.' 0 A + C 1.30:I / '

'"

June July Aug.

520

C\J

11'1380eon

-j.J

~260·rlOJ~~~130HOJ.g8

Figure 4.4.7.5 The effects of physiological age and~te of planting ontuber dry weight.

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Cold70

56

42.p

ib'r-! 28Cl)~

t; 14'0

~ 0.g.p

~ 70.po.p 56G-io~ 42oUl

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Figure 4.4.7.6

Apical

......: t~::loo ,", .....• ~ ,0. :· : ..I :: ".",'~

:"{;.:.-..;'· :'0:;:"., ,..\:;~6

':-.:~{~\'1,' ,-

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24 July

<, 40mm

40-60mm

>60mm

A+C Early Late

14 August

The effects of physiological age and date of plantingon percentages of tuber size'grades out of total tuber weight(nry weight basis). Vertical bars are the SEDs.

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1000 SE~DOO I SEDs

800 80 ...a,/,-

C\J,/

IS r600 60 /till.. /6

8 400 400 IApical 0 EarlyCold 0 Late

200 A+C 20

I SEDs I SEDs

6 508.~ 400

~0 30..!-tQ)

.g. 20+>

* 10

I

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1/ '

60

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o Earlyo Late

June July Aug. July Aug.

Figure 4.4.7.7 The effects of physiological age and date ofplanting on total dry weight (TDW) and percentages of tubersout of TDW.

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•Cl>o>::Cl>eoHCl>ElCl>

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11/

Table 4.4.8.2. The effects of phyad o.Logi cc.L age after 64 days of plantingon, tuber, LAI, 'l'DVJand its components.

Dry weight, g m-2Treat LAI Tuber % tuberment no~m-2 out of

Tuber Leaf AGS UGP TD\r1 TDW

4D 1.03 0.9 0.01 42.3 27.5 14.0 83.8 0.01

24D 1.26 13.5 1.9 52.4 33.5 15.6 103.4 1.8

48D 1.36 31.7 7.6 59.9 38.5 17.3 123.3 6.2

80D 1.80 23.4 10.6 73.7 47.1 21.4 152.7 5.5

128:0 1.79 35.5 21.9 75.7 47.9 21.0 166.5 11.2

184D 1.70 20.9 10.9 '71.1 45.8 18~4 146.2 6.0

232D 2.09 40.0 30.8 87.8 8'7.4 23.4 199.5 13.8

280D 1.65 26.9 11.5 72.6 4'7.1 21.4 152.5 6.0

352D 1.62 21.1 10.6 68.5 42.3 22.3 143.7 6.8

920D 2.25 52.3 4'7.6 95.0 59.3 25.8 227.7 20.4

SED 0.237 12.06 9.38 -9.67 '/.25 2.95 25.02 4.34

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11~;

4.5 DlSCUS:Jlm:

4.5.1 Sorout growth during storage

Similar results were obtained over three years for sprout growth

during storage. Total sprout length per tuber as well as growth of

the indi vi dua'l sprout incre use d with i.ncr-ease in il1i t ial, tuber weight

and there wc's a s i.gni f i can t linear r eLa t ioneh i.p between total sprout

length per tuber and initial tuber weiBht. This may reflect the avail-

ability of substntes (~~orris, 1966, 1967j ;.,rurr, 1978;).). Total sprout

olength per tuber mer-eased wi th Lncr eaee in day degrees above 4 C after

dormancy break for a similar type of treatment (sprout growth rate was

different for different types of treatments e.g. ap ica.l , multi, fast

and slO\,,). '."hen lines of cumuLc t ive total sprout length per tuber as

40 'a function of cumulative day degrees above C from dor-mancy break over

three years for :.Apicdl tre,'; tmen t from the experiments: F1 (both varieties) ;

F2jF3 were compared, they differ in intercept iilld slope which was

related to initial tuber weight. Thus for these .lines multiple re-

gression was done in which day degrees above 4°c from dormancy bre ak

accounted for 70;; of the var Lance (significant, P = 0.001) in total

sprout Length per tuber .irid when tuber ve i.ght W:J.S included the variance

accounted for increased to 80% ilnd this increase due to initial tuber

weight was significant (p = 0.001). So the tobl sprout length may

be represented by the equation,

SPL = 2.25( "!: 2.08) + 0.31( t. 0.002)DD + 0.118( t, 0.023)T\O/

-1'\<Jhere, SPL = Total sprout length, mmtuber ,DD = day degrees above

4°C from dor-mancy break and T;i/ = Lni t i al tuber weight , g and residual

standard deviatioI,1 = 4.58 and residu.J.l degrees of freedom = 53.

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When data from the experiment F3 w~s also included in the multiple

regression the var i auce .iccounted for decz-eased from 80},j to 7Qb; perhaps

the dormancy of this seed lot hud already broken be f'oz-eit was re-

ceived, for this seed h ad many d.unaged sprouts which Lnc reased the

number of growing sprouts arid even tu.s.l.Ly the total spr ou t length per

tuber. Increi:J.sein sprout length with iEcrease in day degrees, when

stored at different temperatures during storage immediately after

dormancy break has been reported by se veral.worker-a, Rawi, 1981; Ali,

1979; Wurr, 1978b. When tubers were under ideal conditions for

sprouting Lnmedi a'teLy after dorrn..ncy break, apical dominance established

within the sprout ~opuliltion, l~rger sprouts inhibiting the smaller

ones as reported by Goodwin (196}), for the variety Arran Pilot.

Storing tubers in cold (3 ± 1oC) stimulated more sprouts to grow, as

reported by \vurr and Allen, 1976, and tobl sprout length per tuber

per day degree increased as reported by Krijthe, ~962 and Wurr, 1978a~

Thom3s and Wurr (1976) reported .1 build up of the gibberellins in

notato tubers following cold storage for 14 days, which may have in-

creased sprout number and thus the total sprout length per tuber.

Al though to t.al,sprout length per tuber Lncr-eused due to greater number

of sprouts following cold storage the extension rate of the longest

sprout was not changed. 'Ivhentubers were stored in the dark extension

'rate 'of the sprouts was increased , this confirms the report of Rawi

(1981 ).

If the physiological st~te of the tuber is to be judged from the

state of the sprouts present then tuber weight must be taken into

consideration. If tubers were stored ~t conditions ideal for sprout

growth Lmmed i,utely after dor-m.mcy br-eak, then total sprout length or

length of the Longe s t sprou t may be a Good Lndicator of the physiological

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1?()

state of the tuber. But if tubers were stored for different periods

in the cold (3 t 10C) then length of the longest sprout alone appears

to be the par-ameter for judging the phyai.o Logi cal, e ti te of the tuber.

Hean sprout leq~th_\s used by !'~orris (1966), may not be the useful

parameter es oec ia.ll.y in the cuse of ap i ca l.Ly sprouted tubers where

some sprouts stoPged growing.

Emergence

In these exoe r-Lment s the Longe s t sprout in the case of apical

trea.tment wus 7 - 16mmlonger than that of multi and in three out of

the four cases, apic:ll emerged Cl day before multi but this difference

was not s i.gni.f'dcan t and may not be of much pr-act icul, importance,

e.imiLar but slightly different results have been reported by other

workers e.g. 'vlurr and Allen (19'('6), could not find differences in time

to emerg~r:..ce in seed lots sprouted either from 15th Se;:>tember, 15th

November or 15th .Innuar y unti L planting. Ali (1979) found a difference

of 1 - 4 days in appe ar ance of 50,i£ pl:111ts in various experiments where

difference in length of the Longe s t sprout v.rr i ed from 6 - 3Omm.

Younger (1975) found 3 difference of 4 days in one ye~r (planted on

23rd April) .md 1 day Ln ano the r year (pL:mted on 3rd !.1ay) between

two treatments, LS (sprouted for 12 - 13 weeks at 120C before planting)

and SS·( sprouted for 18 daYB ut 120C before pl~ulting), in time to

reach 50% emergence. Rawi (1981), reported that 50X of plants appeared

l~ to 5 days earlier in phye i oLog ic.d.Ly old seed (longest sprout over

100mm) compared to phys io.Logi.cal.Ly young seed (longest sprout 32mmin

one exueriment 3.nd 62mmin another). However tubers with such long

sprouts may be .imnr-act i cal, to be used for commerciall)l.:mting.

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121

Harvesting before emergence showed th., t af te r- plan tin..;, w i th the

availability of moisture and nutrients, the ex te ns ion rate of sprouts

increased many fold and i t 'IJc:~S 2.8mm d;;;.y-1 sprout-1 between 7 to 14

da.ys of p.Lan tLng (Experiment F3, 1980) arid this may have further in-

creased before emergence with increase in soil temperdture (Headford,

1962; Bremner and Radley, 1966; Borah, 1959). Furthermore it is not

only the longest sprout which hue to emerge and in the case of apical

sprouting some branches had already ini t isted in the store on the

main sprouts, and these developed as branch stems. Branch stems

usually emerged later than the main stems. In one experiment (F2)

where apical h ad a higher proportion of br-anch stems compar-ed to

other experiments 5Qj emergence \':,:,-s one day later th.m the multi treat-

mente Thus gr-eate r differences in emergence due to differences in the

length of longest sprouts mav not be expected especially when planting

is done late e.g. after the middle of April.

Experiment F6, which was taken for emergence showed that when at

oleast 80 day degrees above 4 C wer-e given just before pL.mting emergence

was only delayed for a day or so and the difference was not signifi-

cant but less them 80 day degrees above 4°c did sugn i f'Lcan t.Ly delay

the emergence. Depending upon the time of planting in different exp-

eriments and different treatments in the same experiment, cold emerged

'4 - 14 days later than the seed which had at least 80 day degrees above

4°C before p.lan t ing , Similar results wer-e reported by Younger, (1975).

It ';Ias seen that about 50 day degrees above 4°C wei:e required to have

sprouts of about 2.3mm in the store. About 5QS to 60;;j of the sprouts

were either removed or damaged in mech.m i cal. p.lant i.ng from seed lots

which were either stored ut 12°C for, 12 - 13 weeks or 10 days, before

planting (Younger" ~975). If further tuber development is not affected

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122

(discussed later, 4.5.3), then tubers given about 50 d.cy degrees before

s tor-age may be of practiced import:mce for commercia.l p.l an ting where

mechanical hand l i.ng is nece saar-y , ..:tlthough emergence m:.lybe delayed

for about 2 to 3 days , Results may v::'..ryfrom one varie ty to another

as in Experiment F1, where two vo.rieties had little difference in

sprout length but Record started emerging ten days L:l.ter than Pentland

Crown. Similar results were obto.ined in 1980, when f'ew tubers with

similar .physiolot:;ic.:U state of t\>10var i.ei ties were planted to see

the effect on emergence. Th3 period oetween plantinG :J.nd emergence

reduces as p.l an t i ng is deLuyed (Bremner and Radley, 1966). The variety

Pen tland Crown took 24 days in 1979 (plan ted on 1st lfi;a.y)and 33 to 34

days in 1980 and 1981 (planted on 15th or 16th April) from planting

to reach 5aJ~ emergence. Simili~r results were found by Younger (1975).

These resul ts were confirmed by da te of plan ting experiment (F5)

in 1981.

Tuber growth and development

T\ew tubers may form on mother tubers during lonGer storage period

in the dark without foliage being produced (ClaveI', 1975; van Staden

and Dimalla, 1977). Van Loon and Houwing (1981) reported reduction in

'incubation period (peiod between the appearance of the first sprout

on the tuber and formation of tu'gers on the snrout when stored in

darkness (ClaveI', 1951) ) by storing at 120C ocompared to 4 C. Rawi

(1981), reported f a.i.Lur-e in emergence due to 'little po t a to ' disorder

in physiologically very old seed. In the present experiments no diff-

erence between tuber initiation (TI) or La te r tuber growth was found

among apical, multi" f'us t and slow t.reu tmen ts of various experiments,

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1?3

but cold (seed stored a t 4°C ur.t iL .. d."y before p.Lunti.ug ) , did deLay

the TI even whe~ considered from emer~ence. This confirms the report

of Raquf (19?9). LAI 1 t t.he t i.me of 'I'I \VU.!3 higher i:l cold compar-ed

to apical and r.u.lt i '-1.S found by Raquf , (1979). 'I'hom.csand \tJurr (1976),

reported an il1cre:J.se in sibberellins following cold storage for 14 days.

\\furr et i.J~., (1980), found hi.gher- concentrations of cytokinin and

gibberellins in 'little potato' comnared to normal tubers and further

they s ta ted that 'little potato' initiation may huve occurred at low

gi.bber-eLl Lns levels but tha t gibbez-el l i.n v.c t ivi ty subsequently increased

vii th tuber growth. It may be t.hut during storage at 120C concentrations

of cyto~inins in the sprouts may have increased while in cold stored

tubers concentrations of gibberellins were increased, so the ratio of

cytokinins and unknown tuber i zc.t i.on stimulus to gibberel1ins may have

been more in sprouted tubers (o.pic;,l or mult i ) compar-ed wi th unsprouted

(cold) at the time of emergence. 'I'h i s resulted in dif'f'ez-ence s in time

to TI (Chapter 2.5). A: though there wer-e differences in phy s i.o.l.ogi ca.L

age be tween ap i cuL and multi t r-e.rtment.s , TI ·.vasnot ;,ffected. Possibly

the ratio of cytokinins ~ld the unknown tuberization stimulus to

gibberelli1".s increase at higher rates immediately on trc:nsfer to warm

conditions (12oC) f'ol.Low.i.ngcold s torsge and after that it increases

. at a s l.ower rate, thus differences may not have been big enough between

ap ical. and multi to affect the '1'1. It appears that cer tai,n aspects of

tuberization are var Leta.l, char-ac ter-Ls t i cs as Record took less time to

initiate tubers and further it ctllocated more assimilates to the tubers,

compared to Pen t l and Cr-own;

In the apicsl treatment L< slightly higher per-cent.age of assimilates

were allocated to the tubers compar-ed to the multi tr-eatrnenta in Exp-

eriments F1 and F3,but not .ir. Exper i.ment F4, although this was not

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significan t , This effec t .rooenr-s to oe due to compet i tion between

stems rather thanphysiologic:.::.l 1:,ge, :'lS total stem number were higher

in multi of Expe r-Emer; ts F"l and F3 «nd de cr-e..lse in sp·.lcings between

plan ts did slightly de cr-ease the pe r cer t,)ge of '~lssirnilates a.lLocat ed

to the tubers. ro difference .in pe r-cen t age of tuber by wei.ght out of

total dry weigh t vIas found be tween cold .ind :tpic',_l or mul ti when COTl-

sidered from thw dite of TI. So it apDeurs that once the TI had occurred

the effect of phyai.o.Logi co.l age di s.ippeur-ed and major f ac to.r affecting

the tuber growth may then h.ive been the environment. An effect of

photoperiod could not be detected i.l.S the difference in emergence between

early ~U1dLate planting of Experiment F5 was only four days and day

length, dur Lng mi d-Nay , when p.Lant s wer-e emerging, was increasing at

the rate of 20 minutes per week. It iG accepted that there is a balance

be tween gr-owth of tubers and rest of the pl.:..tnt anything which favours

the growth of one will retard the gr-owth of others Oloorby,. 1978;

Ivins and Bremner, 1965). II: 19'79 due to \'nter-strcGG h.iu lrn growth was

reduced and tLis resulted in eJ.lloco.tion of ,cl higher percentage of

assimilutes to the tubers compared to very we.:t ye'J.r of 1980. For

example after 36 and 47 days of 50% emergence percentage of tuber dry

weight out of tot al, dry weigh t Wd.S 36 and 53% in 1979; (average of apical

. and multi for Pentland Crown only) 28 and 47/~ in 1980 (Experiment F2,

aver-age of apic.:.U and mul ti); 30 and 48'){,in 1981 (Experiment F5, for

ap i.cal , average of two da te s of planting) respectively. Although a

higher percentage of assimilates was ulloCdted to tubers in 1979 the

overall growth of the cr-op was very much reduced (4.5.4 q, v.) which

reduced the bulking rate per unit ar ea , For examp.Le af te r- 36, 47, 57,

-269 and 81 days from 50}{ emergence tuber weight gm was 107, 227, 362,

432 and 525 in 1979·,(mean of apical and mul ti for PentLand Crovm); 154,

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125

349, 557, 795 and 1091 in 1980 (mean of apical and multi for Exp. F1)

and 88, 234, 359, 503 :md 637 for 1981 (for apical, mean of two plant-

ing dates, Experiment F5) respectively. These figures appear to be

related to rainfall d~ta for three years (Figs.3.3.1 and 4.4.7.2).

Similar resul ts were found by Chowdhur-y, 1980. Llewelyn (1967) in-

creased tuber bulking by irrigation. HcDermott and Ivins (1955),

found a linear relationship between total tuber yield and the May -

September rainfall over eight years (1947-54). There was increase in

yield of 1.4T/ha for each cm of rainfall, which Harris (1978), states

is similar to yield responses found in irrigation experiments in

Britain. Hixing of two type of seed tubers did not affect the bulking

rate as their yield was not significantly different from that expected

i.e. average of two types of seeds when grown alone. But sprouted seed

had the advantage over cold early in the season as found by Chowdhury

(1980). In general senescence (4.5.4 q.v.) was not affected by any

trentments w ithi,n 'CltX1)eriment,50 f inal. yield did not differ due to

treatments, but final yields of 1979 were much lower than 1980, as the

crop was affected by drought in 1979. Significant linear relationship

was found between final tuber yield and leaf area duration (LAD) as

reported by several workers (e.g. Gunasena and Harris, 1968, 1969, 1971).

Several workers (e.g. Gunasena and Harris, 1968, Bremner and Radley,

1966; Bremner and TahL.t,1966) have reported improvement in the relation-

ship between LAD and tuber yield, when leaf areCl.indices above three

were assumed as three, but this was not the case in the present in-

vestigation even assuming LAI over 4.0 as 4.0 did not give any relation-

ship with tuber. Probably light interception increases with increase

in LAI over 3 and further efficiency of the canopy was increasing with

increase in LAI (4.5.4 q.v.). In f'act Lu ter- Gun;).senaand Harris (1971)

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126

also could not improve relationship be tween LADand tuber yield by

assuming leaf ar-ea indices over 3 as 3 or over 4 as 4 or over 4.5 as

4.5. A Ld.near' relutionship be tween tuber number cl.ndtot a.l stem number

was found as reported by Allen, 1972jlOosey, 1962j \'!urr, 1974. Since

stem number does not toke ::..nY:.l.ccount of the s i ze of different types

of stems the relationship between LADuccumul.at ed over & period of

30 days from the date of TI accounted for more var-Lance , This confirms

the mechanism discussed in Chupte r 3.5 th a t tuber numbers which develop

depend upon the assimilates available at the time of TI LU1dCl. period

after that. Sizes of the tubers depend upon the tot3.l assimilates

available for their growth from TI until harvesting. Since LADwas

much greater in 1980, there were higher percentages of bigger sized

tubers in 1980 compared to 1979. The number- of axillary branches

(AB) decreased w.i. th an increase in stem numbers per uni t area

(Ifenkwe, 1975), which in the oresent investigation were related to

leaf area duratior~ (LAD) for a period of 30 days from emergence (i.e.

speed of ground cover). Tuber numbers increased with decreilse in

spacing so their average size decr-eased (Ifenkwe, 1975) thus the pro-

portion of bigger sized tubers may be related to the AB. In the

present investigation tuber yields of relatively bigger sized tubers

were significantly_ linearly related to the LADcontributed by the

axillary branches.

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1?,?

General cro,) groi.,rth

Stem number is now considered as the unit of population in the

potato crop (Allen and Deem, 19'18). Api ca.l tre atrnen t increased the

proportion of br-anch stems compar-ed "Ii th multi and cold treatments in

all the experiments as earlier found by Younger (1975). Main stems

were much bigger them their counterpartbranch stems. It may be that

because br-anch stems emer-ged later they were suppressed by the main

stems. Hence the type of sprouting treatment given to the seed tubers

must be taken into consideration when calculating plant populations.

iv'ater stress may inhibit the formation of new Leuves (Nunns and Pearson,

1974; Zaag and Burton, 19'18) and so may reduce the LAI (Boyer, 1976;

Hunns and Pearson, 1974). II' the present experiments due to more rain

fall in 1980, stems were much longer them those in 1979 and leaf numbers

were increased. This resul ted in higher LAI values when compared with

1979 ill1d1981. Increase in LAI or ground cover with irrigation has

been reported by Llewelyn, 196'1; Hohindra.,. 1975. In the absence of

any appar'en t water stress, higher LAI in 1980 resul ted in higher rad-

iation interception and thus total crop growth was higher in 1980 when

compared \vith the other two years. It muy become more clear if light

interception und crop eVCJ.pora.tion are taken into consider.::ttion. In

general different tre'J.tments within a purticular year did not differ

in their photosynthetic efficiei1cy or evaporation rates thus the results

are discussed in general for two different years.

The fraction of r-adi.at ion intercepted by a canopy depends mainly

on LAI (Shibles and 'weber, 1966; Horie and Udayawa, 1970), and trans-

mission of Lndi v.i.du.d. leaves and their geometrical urrangements. Light

interception (LI) i·li'creCl.sedlinearly until LAI was about 2.25 and

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12B

thereafter L1 Lncz-e aaed at ,1 dim .n iah ing r! te, due to in ter-leaf shading.

Furthermore, Lea f ang.le 1<1::-\yvd.ry w i th (A.ll Lr.cr-ease in Lea f size and so

may affect the geometr-Lcal, :.:l.rr_Elgrnentl'li thin the CJ.}lO?Y. IE the present

investigation these cornbi r.e d effects were de te c ted cis a ch.inge in k as

LAI increased. The efficiency of conversion of light to dry matter

can be estimated from the Ldne.rr relationship betwee n accumulated total

dry weight and ac cumu'la te d .inte r ceo ted T:).diation (Littleton et &1.,

1979; !Jlilford et al., 1980)..vhen forced through the origin (Smillie,

1966), the r-eI.ationships found here .indi ca ted that for every 1.OMJ of

PAR intercepted, the crop produced 3.42g of dry weigh t ill 1980 and 2.49g

in 1979. 'I'he 1980 results are compar-able with results reported for

other crops, for exampl e ; 3.2g;MJ PAR for barley and whea t grown at

Sutton Bonington and Rothamsted (G:.illagher and Biscoe, 1978); 3.5g,MiJ PAR

for sugarbeet (Biscoe, and Gallagher, 1977). The conversion efficiency

in 1979 was much lower than tha t in 1980, probably because 1979 was a

dry year. Reduction in photosynthetic efficiency due to water stress

has been reported for potato (Shekhar- and Iritani, 1979; Chapman and

Loomis, 1953; ;';oorby_ et ell., 19'75), barley (Legg et· ..tl., 1979; Biscoe

et al., 1975j Gallagher and Biscoe, 1978), maize (Verdun and Loomis,

1944), appl e (Schneider and Childers, 1941), soybeans (Schibles and

~veber, 1966) and whea t (Gall:.igher '..ind Biscoe, 1978). A reduction in

wate r supply will frequently cause s tom.itu.l closure (Hoorby et al.,

1975) and thus increase the resistance to C02 uptake. Also decreased

crop evaporation may result in higher respiration rates, associated withhigher leaf temperatures and consequently ne t photosynthesis may be

decreased (Lomas et al., 1972), (evd.poration mesurements for the presentinvestiga.tion are discussed later). Another factor which may have

affected the photosynthetic efficiency of the canopy is the irradiance

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129

level in relutioll to the light sdtur~tion point of individual leaves.

Changes in LAl result .in differences in the degree of ill ter-leaf

shading and consequently alter the irradiance level required to saturate

the leaf cano:;JY. Individuul leaves may be light satur.J.tedat irradiance-2 8 -2 -1of 100w m for whea t (}brsh.:llarid .3iscoeI 19'77)or 50 UEH S for

potatoes (Ku et ,11.,1977). Howe ver , although incident irradiance may-2 -1reach 2000 UEl'] S ,in summer, the canopy as a whoLe is seldom light

saturated. In 1980, in the absence of any apparent water stress, the

pbotosynthetic efficiency of the cdnopy iLcreased slightly with in-

creasing LAI, up to LAl values of around 5 (Figs.4.4.6.5j 4.4.3.16 ;

4.4.3.18; 4.4.3.19). Further evidence for increased photosynthetic

efficiency of the canopy with increasing LAl comes from the relation-

ship between CGR and mean leaf ar-ea index (L) (Fig.4.5.1). Up to LAl

values of 2.5 - 3 the relationship between CGR and L was similar to

that between light interception and LAL However at v<.:iluesof LAI

gr-eat.er than 3 tot.i l Light in terception increased at u slower rate than

CGR i~e. the conversion of light to dry matter was more efficient per

unit area when LAl was higher than 3. Puckeridge and Ratkowsky (1971)

reported a similar effect in wheut u.lthough the values of LAI were

higher due to the characteristics of a grass type canopy.

At LAl values of 4 or more around 95%of the incoming radiation

was intercepted and it muy be eXlJected thdt for some of the lower leaves

the rate of respiration would exceed gross photosynthesis. However at-2these vulues of LAl the total dry weight of the CiJ.IlOpywas over 1000g m

and:consequently the contribution to total cunopy respiration, by the

minor proportion of Lower Leaves, woul.d have been smalL, As has been

shown in other crops (e.g. Ga.lLagher and Biscoe, 1978), crop growth

rate was found to be,linearly related to PAR intercepted by the leaf

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canopy. The relationship was clearly influenced by water stress as in

1979. Although the efficiency of light conversion to total dry matter

was much reduced by water stress in 1979 the percentages of tuber dry

matter out of total dry we igh t wer-e increased (4.5.3 q.v.). A linear

relationship between tuber dry weiGht and total PAR intercepted over

the whole season was found for both years data as reported by Scholte

Ubbing (1959).

Soil moisture studies of the present investigation showed that in

1979 due to water stress and lower LAI crop evaporation viasmuch lower

than the potential evapor-at i.on (Penman, 1956). 'I'he se differences were

grei:\ter.than the differences detected for pei:l.s(Dawkins, pers. comn.)

for the same year but the peas were planted earlier than the potatoes

and so they were better established than potatoes by the commencement

of the dry spell. Fulton (1970) described potatoes as much more

ser...sitive to water stress than maize or tomatoes. Burrow (1969) demon-

strated that the rati,oof actual to po ten t ial, evapor-at ion fell more

rapidly in potatoes compar-ed to sugarbeet. Shepher-d (1972) reported

a greater reduction in crop evaporation of potntoes compared to mixed

crops of grass and clover. He related this to the greater sensitivity

of potato leaf diffusion resistance to'etdecrease in leaf water potential.

Fuehring et al., (1966) reported considerable reduction in crop evapor-

Cition if irrigation was not given at 75% available soil moisture.

Potatoes are consider-ed to have shal.Low root systems compared to other

crops e.g. sweet corn, tomato, sugarbeet and barley (Corey and Blake,

1953; Durrant et aI., 19'73). In the present investigation roots extracted

water from a depth of 90cm in one year and 80cm in another i.e. in the

orders found by French, et al., 1972; Durrant et al., 1973. Ark;Ley(1963)

showed that there WaB a linear relationship between the amount of dry

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131

matter produced and the umoun t of w:~ter evnpor-ated, Penman (1963)

related tuber bulking to the adjusted potenti~l evaporation. In the

present investigation crop growth rate VIas linearly related to the crop

evaporation. Although there was a difference betweeriyears, wi thin any

year, physiological age or storing tubers in the durk before planting

did not affect the overall to ta.l gr-owth of the crop when considered

from the date of 5(},l;emergence. However the proportion contributed by

different types of stems WelS different for different physiologically

aged tubers" as apical,treatment increased the number of branch stems.

When total stem number were increased either by decreasing spacing or

using bigger seed, LAI and total dry weight were increased early in

the season, and this resulted in more tubers being initiated. The

average tuber size was reduced as found by Ifenkwe, 1975. General

growth of the mixed plots, where two types of seeds (i.e. sprouted and

unsprouted) were mixed within rows was not different from expected

(i.e.').verage of sprouted (apical or multi etc,) and unsprouted (cold)

when gr-own alone). Sprouted occupied more space than allocated to it,

this confirms the report of Chowdhury (1980) but both treatments sene seed

at the same time.

One way of increasing the interception of total radiation is by

increasing the longevity of the crop. In 1979 senescence of the crop

was not affected by the physiological age or spacing but the variety

Record senesced after 138 days of p.lanti.ngand PentLand Crown after 160

days of planting. In the case of Pentland Crown it \'JaS observed that

small seed (34g) emerged later and senesced later than bigger seed

(62 or 105g). In 1980 there was no difference in senescence among the

various treatments of Experiments F2 and F3j all senesced after 162

days of p.Lant i.ng, 'I'r-e atment 025 of Exce rimcn t F4 sencsccd 16 days

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132

later than other treatments of that expe r i.ment .ind 12 days later than

the Experiment F2 or F3. Differences in LAI due to different treat-

ments of Experiment F5 (1981) disc.tppe'J.red from the middle of July and

ligh t interception measurements showed that about 93% of the incoming

radiation \VUsbeing intercepted on 30th. September and 90X on 13th.

October, w i th no difference between treatments. It now appears that

senescence in 1979 1:J:J.S affected due to water stress. As large seed

emerged first it may have been affected more by the water stress than

the later emerging small seed. This hypothesis agrees \'lith Bagley

(1971) who foung than an eu.rly developing crop suffered more from the

drought ·in July than the less advanced crop. Younger (1975) reported

that cold treated seed emerged later and senesced later than the sprouted

seed. Sene ac ing in 1980 was affected by the very humid weuther- as stems

were over a metre long, lodging occurred and rotting of stem tissues

was observed from the middle of August onwar-ds, Another f ac to.r could

be the depletion of nutrients especially the IJ (Ivins, 1963; Ivins and

Bremner, 1965; Gunasena and Harris, 1969, 1971), as crop grew at a very

fast rate. Due to higher stem numbers total dry wei.ght of all treat-

ments of Experiment F4 but S25 was higher than the treatments of Experi-

ments F2 or F3, so they might have depleted the soil before others.

Another factor vrhich may have affectc-d the senescence is the transmission

of wavelengths above 700nm (Holmes .md Snith , 197'7; Scott et al., 1968),

which may have affected the physiological status of the plants as LAI

was very high in that year. Another evidence for this come from

Chapter 3, where the pp333 treated plot in 1980 had a close canopy

compared to other treatments and this F.iayhave af'f'e c te d the transmission

and it senesced before others. It caL not be the temperature as some

individual guard pLHltS were aee n green for itt least ten days after

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133

the crop had senesced. ;JO\"I, since 1981 has been the medium year for

rainfall and LAl did no\;reach above 3.5 and growth rate has been much

Lowe r' than 1980, thus soil may not have been depleted for nutrients

and as there was no severe drouGht like 1979 the crop did not senesce

until the middle of October.

For delaying leaf senescence in :...we t year, application of nitrogen

later in the season may be helpful (Gunase n« and Harris, 1969, 1971).

Later application of N may slightly affect the bulking r-ate, but if

L~r of over 3 or so is maintained during September and the middle of

October could be very useful as due to short days most of the assimilates

produced may be used for tuber growth only. In a dry year like 1979,

if irrigation is given in such a way that crop does not suffer from

drought and LAr stays around 4.0, may be helpful in increasing the

duration of the crop.

Any of these techniques to ext.end leaf persistence by delaying

leaf senescence could be frustrated by blight disease Ci.ndof course

blight control in itself extends leCJ.fpersistence und the period of

tuber bulking.

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4U

3S

+ + + 2CGR = 2.03 (-0.98) + 8.09 (-0.89) L - 0.67 (-0.16) L

% variance accounted for 76.6residual standard deviation = 4.65

30 0 Experiment F21::. Experiment F30 Experiment F4 0

[:;

ne:: °0 0(_,.._; -------_...-_..----

/-

/'/' 0/'

r< / [:;I

>0 / 0III /'0 2JN CJIE

//

/

0' /0..~o /U /0

15 // [:;

[:;

10uj/

5

o~

O-t-----r-------.------,----------,------- ---,-----,------,2 J 4 5 5-1 J 1

Mean leaf area index (L)Figure 4.5.1. The relationship between crop growth rate (CGR) and mean

leaf area index for all experiments in 1980. Each data point is anav~rage fdr number of plots in that experiment.

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134

SUMMARY AND CONCLUSIONS

In the majority of the field experiments, final tuber yields

were not significantly different over a wide range of treatments

which included (a) different physiological ages (b) different

sprout growth rates at planting time (c) different stem populations

obtained by adjusting plant spacing and seed tuber sizes

(d) a mixture of seed tubers of different physiological ages

or even (e) the two different varieties. In 1979 drought in

early summer affected all treatments but there was some recovery

when rain fell later, but because of this all yields were lower

than in 1980. The photosynthetic conversion efficiency of the

canopy (g dry weight MJ-l PAR intercepted) was 2.49 in 1979

compared with 3.42 in 1980 when there was no apparent water

~tress •

Final yields are multiples of tuber numbers and mean tuber

weights. and these two components, together with stem numbers,

were significantly different with different plant spacings,

seed tuber sizes and storage treatments. Record produced

mqre tubers than Pentland Crown. The relationship between seed

tuber numbers and stem numbers was linear and significant.

In treatments which gave higher stem numbers in the crop

L.A.I. and general 'crop growth rate were higher early in the

season but other treatments caught up later. PAR interception

increased linearly with increases in L.A.I. up to LAI ~ 2.25

when over 70% of the incoming radiation was intercepted.

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135

Above LAI = 2.25 the rate of PAR interception slowed down until

LAI = 4.0 when around 95% of the incoming radiation was

intercepted. F~nal tuber yields were significantly rQlated to

the PAR intercepted by the canopy over the whole season.

No treatment other than varietal differences hastened

leaf senescence which was later than anticipated and this

probably explains why final tuber yields were not significantly

different for several different treatments. Growth analysis

studies showed how different aspects of plant growth were

affected, but in each case it appears that the crops arrived

at similar final yields but by different pathways, i.e. bulking

rates x duration. Growth analysis results also showed that

had the crops been burned off or lifted earlier for final yield

then the different effects of many treatments would"have been

much larger. Hence seed treatments are of vital importance with

early crops and probably second early crops where lifting

occurs before mid-August. However, with maincrops which are

allowed to mature late, by favourable environment, absence

o~ blight and by production treatments such as irrigation or

higher N levels, then a great dea~ of catching up takes place

a'ndat final harvest there are not likely to be great differences

in yield from a range of seed treatments. The major effects

are likely to be on tuber numbers and sizes which can have

implications for quality for various purposes.

Drastic effects on leaf growth resulted from treatment with

a growth regulator pP333. Although this investigation was no

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136

more than observation plots with no replication, it was evident

that leaf area per plant was reduced and higher tuber yields

resulted. It proved possible to plant closer without enhancing

interplant competition and PP333 appeared to increase the

allocation of assimilates to the tubers. The result was

more medium sized tubers. This preliminary trial suggests

that there might well be a future for plant growth regulators

with the -potato crop and gives encouragement for further

investigations to be carried out.

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ALI, S.li.J.i'<:. (1979). Effect of tempe r-atur e dur ing SprOUCl!1g on gr-owthand yield of t!1C m.ii n cr op ~)OLitO V .r i.e ty Desiree. l·i.Se. thesis.Uni versi ty Col Lege 0 fd a.i es , Aberys h:y tho

ALL1~I:,E.J. (19?2). The c f'f'oc t of r ow w i dt.h on the yield of threepotato vrr Leti.e a , J our nal of l'\.~~ricul tur a.l Sciecce, Cambridge, 79,315-321.

/l.LLEI~,s.a, (19?8)~ ?J:'Lt density. r.. Fi'he Po t.i to Cr-op, The scientificbas i s for imnrovement' (Ed. ILrris, i).H.), np 2'('8-326. Ch.rornan andHull, Lor.dori,

ALL;~r.:,:':;.J.ud;]~;}\l:, J.:. (1978). Fictors influencing the rel0.tiol1-sh i.» between tuber yields ind popu.lo.t ion , Pot..:to Research, ~, 51.

ALLEIJ,~.J., 3;:;ji;" J.::., GEH'FI'j_'Il, r:.L., O'CRI1:;L, P.J. (1979). Effectsof length of spr-ou t i.ng period on grm-/th and yield of con tr as t inge:J.rly pot~to vurieties. Journul of A3ricultur~1 Science,Cumbridge , 92, 151-163.

ALLEri, E.J.'J.l}d SCVr'1:', 1-:.::. (1980). An ..:!l::llysis of grm':th of the potatocrop. J our-nul, of Agricul tur a.L Science, Cambridge, 94, 583-606.

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flRTiiUR, .i.n., GU'l'lIRI.e;,J.~) •.. .nd [;E'3£1,L, J.I·I. (1930).ar-t i f i.c i al. cl irna te s on the grO\-/th and chemi co lAmerican .Iour-na'L of Bot uny , 17, 416-482.

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BODLAENDER,K.:j.A. and EiillIl:U.s, J. (1969). The influence of the mothertuber on Grovlth and tube r i zu t i.on of [Job. toes. l.e the r-Landa Journalof A(iric. Science, 2Z, 300-30S.

OODLAiJJDER,Ko3.A. arid NAnEUS, J. (1981). Aspects and preliminaryresults of r-ese ar ch OIl physiologic.:.::_~geing. Pl an t growth undercontrolled conditio!1s. Proceedings of the 8th Triennial Conferenceof the Eur-opean Aaaoc i.a tion for Potu to Research, pp '70-71.

BOOTH,A. (1963). The role of Growth subs tcnce s in the development ofstolons. In 'The gr-owth of the po ta to ' (Eds. Ivins, J.D. andfhlthor:Je, F.L.). ~)l) 99-113. 3utten:orths, London.

BORAH, t·1.N. (1959). The effect of Ligh t Ln tens i ty , length of day andt.empe r at ur e on gr-owth .ind tuber f'orrna t i or; in the po ta to , Ph.D.thesis, Uni versi ty of Not.t ingh::un.

BORAH,11.!:. and :.;rL1,!lOEP;~,F.L. (1962). Growth of the potato as in-fluenced by ternpe r atur-e , Lnd i an Journill of Pl an t Physiology, ~,53-'72.

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APPENDICES

Appendix A

John Innes potting compost number was prepared by mixing loam:

peat: Grit; 7:3:2 (volume basis). 372g of J.I.B (5% N,

7.2% P205 soluble, 1% P205 insoluble and 10% K20) fertilizer

and 70g of chalk was added per 100Kg of mixed compost.

Appendix B

Seed used for experiments: GR1 (growth room); F1; PP333 trial

1979, was obtained from UCW, Aberystwyth, where it was grown

at Rhayader, Powys, from Scottish FS3 (Pentland Crown) and

FS2 (Record) stocks. It was planted on 10 May, defoliated on

4 August and harvested on 5 September

Appendix C

Seed used for experiments: GR2 (growth room); GH1 (glasshouse);

F2 was also obtained from UCW, Aberystwyth, where it was grown

at Dyfed, near Llanarth from a once-grown Scottish VTSC stock

(Multiplied in 1978 at high altitude seed site near Rhayader,

Powys). It was planted on 22 May, defoliated on 10 August and

harvested on 17 September.

Seed used for experiments: F3; F4; PP333 trial 1980 was grown

at Bunny (University of Nottingham Farm) from Scottish AA1.

It was planted on 7 May, defoliated by the end of August and

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harvested from middle to end of October.

Appendix D

Seed used for experiments F5 and F6 was obtained from uew,Aberystwyth, where it was grown at Dyfed near Llanarth. It was

planted on 17 April, defoliated on 21 July and harvested on

4 September.

Appendix E

Main stem The stems directly originating from the mother

tuber.

Branch stem - The stems originating from the underground stem

i.e. not straight from the mother tuber.

Axillary branches The stems originating from the leaf-

axis above ground.

Main stolons The stolons originating straight from the stems.

Branch stolons The stolons originating from another stolon

i.e. not straight from the stem.

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