A Quantitative Study of Wate Culture Solutions Review

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A Quantitative Study of Water-Culture SolutionsA Quantitative Chemical and Physiological Study of Nutrient Solutions for Plant Cultures. byW. E. TottinghamReview by: Walter StilesJournal of Ecology, Vol. 2, No. 3 (Sep., 1914), pp. 182-185Published by: British Ecological SocietyStable URL: http://www.jstor.org/stable/2255518 .

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182 Notices of Publicationsof General Bearingsummations; hence forpractical urposes nd for he present, he formermethod, ill now basedsolelyon phenologicalobservations,eemsthus to be placed in closer ogical connectionwiththetemperatureoefficientsf chemical,physical and physiological rocesses han has hitherto eenthe case. (2) The similarity etween he resultsderivedby these-twomethodsof temperatureintegrations, however, nly uperficial nd roughly pproximate;the ratiosof direct ummationto efficiencyummation ange n magnitude, or the mean frostless eason in theUnited States,from minimum f 7-49 to a maximum f 10-44. (3) A rationaland consistent limatic chartrepresents he geographical istribution f these ratio values; on such a chart the marginalregions of the country re frequentlyharacterised y low ratios, nd the twomain mountainsystems Appalachian nd RockyMountains) ppear to control reas of high values. (4) Thereseems to be no doubtthat the ratiohere brought orward epresents climaticdimensionorcharacteristic, hichappearstobe some sort of function f the daily normal emperaturesponwhich thiswhole study as beenbased and of the timedistributionf the temperatureata withinthe periodofthemean frostlesseason.

A QUANTITATIVE STUDY OF WATER-CULTURE SOLUTIONSTottingham, W. E. "A quantitative chemical and physiological study ofnutrient solutions for plant cultures." Physiol. Researches,1, 1914, pp.133-245.

Thispaperon nutrientolutions illbe ofgreatuse to all whoseresearchesecessitateinvestigationsnto he nutritiveolutionfplants r theuseofthewater-cultureethod;andalthoughotprimarilyn ecologicalapert s onewhich as muoh earingn some spects fecology.The paper s dividednto hree arts. Part consistsf a concisendvaluable6siu?nf hework hathas beendoneupto now n water-culture.t is shown hat heuse ofwater-culturemethodsalls nto hree eriods. 1) Theearly eriod,ndingbout 860, stablishedhe deaofthe oilmineralss a source f henutrient aterialfplants. (2) The middle eriod,rom 860to about 900, stablishedhevalueofwater-cultureor ertainlasses fproblemsndevolvedalsostandardutrientolutionsor se n thismethod. Chiefmonghe esults btainedy hemethodn this eriod asthedeterminationf he ssentiallementsor lant utrition.Otherfactswere lso indicated,uchas an optimumotal alt concentration0 30/0, or owerfthesolutions ere hanged requently),lthoughtwasfoundhat he relativemount fdifferentsaltscouldvarywithin atherwide imitswithout roducingnyapparentnj rious ffect.Selectivebsorptionndexcretionfcertainaltswere lso indicated. 3) In thethird ndpresenteriodt is the selectivebsorptionf onsratherhan ompletealts hat s indicated,andso by hismeans he ompositionf henutrientolutionecomesltered. The mportanceofa physiologicalalance etweenhevarious lementsn the nutritiveolution as alsobeenemphasiseduringecent ears.The author inisheshispartof hispaperby pointingutthegreat ariationsn relativeamountsfdifferentubstancesn thevarious tandardutrientolutions,nd so concludeshatit s ustifiableo make a thoroughnvestigationf he elativealues fdifferentroportionsfthe alts mployedn such ulture edia." Theremainderf hepaper onstituteshe eginningof uch piece fwork.In Part I a chemical tudys madeofKnop'ssolution,s a result f which hemostsatisfactoryethodfmakingpa stock olutions described. t is desirablef courseomakeup a solution hichs ofas high concentrations is consistentith tability.Tottinghamsuggestssing wo tock olutions,ne of calcium itratelone, he other ontainingheothernecessaryalts. The latter olutions stable or bout14 days t a temperaturef 23?C. ina concontrationf9?/. or 10 /0 accordings K2HPO4 rKH2PO4 is usedas the source f

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A Quantitative tudy of Water-culture olutions 183phosphorus. The precipitate bove the imit of stability s almostentirely alcium sulphate,notcalcium phosphate s has been supposed.In Part II of the paper we have a record f the experiments hich were carried ut with aviewto determininghe most suitable nutrient olution. Young wheat plants were used, andweregrown or approximatelyhefirst hird f the fullgrowth eriod f wheat. Colourlessflintglass bottlesof 250 c.c. capacity were used, and six plants weregrown n each bottle. Theyremained 4 days n the nutrient olutions,which were changed very hirdday. At the end ofthegrowth eriod heyweredried nd the dryweights f shoots nd roots aken eparately.Preliminary eries of experiments ere made in which Knop's solutionwas used in variousconcentrationsut nwhich a constant atiobetween he various onstituentalts was maintainedthroughout. In the case of each solution ts osmotic pressure s calculated from lectrolyticdissociation ata, but as no account s taken of the influence n the dissociation f one anotherproduced y several altstogethern the same solution, henumbers or he osmotic ressure reonly pproximate.

Tottingham's esults how that concentration as practically o influence n the amount ofdrymatter roduced ver a very onsiderable ange,namelyfrom concentrationf total salts of001 percent.to about one of1-4percent. It is concluded lso thatmonopotassium hosphate sa better alt to use as a sourceof phosphorus han the dihydrogenhosphate, nd it is the formersubstance hat was used in the subsequent eries.In the furtherxperimentshreemain seriesof cultureswere rranged. In thefirsteriesthetotal concentration f salts was 06 0/0,which ies wellwithin he rangeof concentrationsivingoptimum rowth. In the econd eries he concentrationf salts was0-01 /0. This concentration" was selected as havingbeen shownin the preliminaryulturesto lie considerablyelowtheoptimumi," and also because the concentrations probably imilarto that ofmanynatural oilsolutions. In the third eries the concentration,0 ?/, was above the limits of optimum on-centration,nd also presented onditions f concentrationimilar o thosewhich obtain n alkalisoils.In each of theseseries a number f nutrient olutionswere obtainedby varying he relativeproportions f the four salts Ca(NO3)2,KNO3, MgSO4, KH2PO4, but within each series theosmotic pressures f the various solutions remained pproximately onstant. The dryweightswere akenand the resultsplotted n the triangle iagram rrangement sed nphysical hemistryand introducednto botanical onsiderations y Schreiner nd Skinner2*The chief esults f theseexperimentsre as follows:1. Where he otalconcentrationf thenutrientolutionwas 0 6?/.. (a) Where he osmoticratio of MgSO4 was greater han 2-0 =a molecular ratio of 3 23), a verymarked nd curiousCa(N03)2injury o the plants was observed. The tips of the leaves "rolled backward aterally o forma stiff eedle-like tructure and thiswas generally receded yyellowingnd ultimate leachingofthetipof the eaf . Later theyoung eaves became imp and spirally oiled n the middle or

towardsthe base. No injurywas observedwhen the ratio MgSO4 was below 0-25 (= aCa(N03)2 wsblw05(molecular atio of 0 40). The extent f the injury n cultureswhere his ratio was intermediate1 However, he actual differencen dryweightbetween he culture n this concentrationndthose n theoptimum oncentrationss so slight hatthe difference ay quite possibly ie withintheralngefexperimentalrror.2 "Some effects f a harmful rganicsoil constituent." U.S. Dept. of Agric.Butr. f Soils,Bull. 53, 1910; also, " Ratio of phosphate,nitrate nd potassiumon absorption nd growth."Bot. Gaz., 50, 1910, pp. 1-30.3 This formof injurywas observedby the reviewern water-culturesf rye grown n the

spring f 1913. In this case the molecularratio Mag(N04)was as highas 14,while t was onlyslightly n evidence when the ratio was as high as 2-8. The total concentration (075) was howevermuch ower han nTottingham's xperiments0 6). In veryow concentrations01) Tottinghamfound he njury ractically on-existent.



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184 Noticesof Publications of General Bearingbetween -25and 2-0, was presumaablyetermined y the concentration f the other alts in themedium. (b) The bestgrowth f shootswas obtained n solutionshaving the salts in approxi-mately the followingosmotic proportions: 2-5 KH2PO4: 2-5MgSO4: 4 0 CaNO3 1 0 KNO:3correspondingo molecularproportions f 2-66 : 2-97: 2-9 : 1 00, a considerably ifferentarrangementromKnop's. The best solutionfor he growth f rootswas however ound o bequite different,amely ne in whichthe osmoticproportions ere1 00: 0-25 0-25 1 00 (corre-Fpondingo molecularproportions-07: 0 30: 0-18: 1 00)1. (c) The development fthe shootswas notproportionalo that of theroots. Livingston ad found his tobe so in the case of wheatgrowingn soil cultures2, o Tottingham upposes that while in a liquid hnedium the con-centrationsf the variousmoleculesand ions are not greatly nfluenced y the metabolism f theplant, nd considerable ifferencesn root rea mayhave slight nfluence ponthe growth f tops,in soil cultures, n the otherhand,the ctivities f the plant may through xcretion f carbon-di-oxide, etc.)alter heconcentrationfthe soil solution n various rons, nd theextent f therootsystemmay henbecome muchmore mportant actor nthe growth f tops." This explanationdoesnot seemtobe convincing.2. It the cultur s where he otal concentratioinf thesolutionwas 0-01 /0. (a) Both shootsand roots f heplantwere longated nd slender, hisbeing east marked nthe cases ofroots row-ing n solutions owest n concentrationfMgSO4 nd highest n KNO3 oncentration. b) In somecultures yellowingnd dying f the basal leaves occurred, robably ue to deficiencyf nitrate.The lowyields n some cases appear to be due to the same cause. (c) The peculiarmagnesiuminjurywas absent,but thegreatest ieldswereobtained rom ultureswhere heratioof M(g(04)was low,and the eastyieldsfrom ultureswhere he ratio was high. (d) The bestproportionfsalts forthe growth f shoots was considered o be in osmoticproportions, 5 KH2PO4: 2-0MgSO4 : 15 Ca(N03)2 : 10 KNO3 correspondingo molecularproportions f 0 34: 2-00 100:1.00). In the case of rootsthe best osmotic roportions ere06 : 02 : 0 2 : 10 (colrespondingto molecular roportionsf 0-41 0-20 0-13 1 00). (e) As in thepreceding eries he yieldofshootsboreno relation o theyieldofroots.3. In the ultureswhere hetotal oncentrationf salts was 2 00/0. (a) Root development asadverselyffectedhroughoutheseries,butchiefly heretherewas an excess of any one of thenutrient alts. An excess ofhydrogen, alcium or magnesium onsappeared o be most njurious.(b) A high proportion fcalcium appeared ocause wilting f the eaves and ultimately hedeathof the plant. (c) In some culturesthere appeared a yellowing f the basal leaves followedbydeath. This mighthavebeen duetoa lowproportionfmagnesium ulphate, utwas notclearlyso. (d) In other ultures he tissuebetween he eaf veins became bleached. This was probablycaused by the high concentration ombinedwith certain unfavourable roportions f salts.(e) Magnesium njury ccurred s in the first eriesbutno simplerelationbetweenmagnesiumandcalciumcouldbe obtainedn this case. It is clear that theantagonism etween alcium ndmagnesium s much influenced y concentration. (f) The best proportion f salts forshootdevelopmentwas found to be *22 KH2PO4 : *67MgSO4 : 33 Ca(NO3)2: 100 KNO3 in osmoticproportionscorrespondingo molecularproportions-23: 081 : 0-25: 1 00). In the case ofrootsthe best osmoticproportions as considered o be 04 : 1-8 : 0-8: 10 (correspondingomolecular roportionsf0-41 2-20 0 59 : 1-0)3. (g) There was notmuch correlationetweengrowth f roots and shoots in this series. (h) A greatdecreasein the ratio of water oss bytranspirationotheyieldof shootaccompanied xcessivemagnesium njury. In the first eriesno such correlation as obvious, nd the decrease n water oss in this series s presumablyueto thereduced eafsurface n themuch njuredplants. The transpiration erunit ofdryweight

1 These appearto be thecorrect umbers, nd are so givenbyTottinghamn thesummary fthepaper. Those actuallygiven t thisstage ofhis paperarequitedifferent.2 " Note on the relationbetween he growth f roots and of tops n wheat." Bot. Gaz. 41,1906, pp. 139-143.1 The numbersgivenby Tottingham t this stage of his paper re quitedifferent.The onesquotedhere are takenfrom is summarynd are presumablyhe correct nes.



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A Quantitative tudy of Water-cultureolutions 185was also much ess than n the second series, resultwhich was also found, hough o a muchless degree, n the cultures n weakest oncentrationf nutrient olution.In conclusion t would eem necessary o call attention o two criticismswhich may bemadeon the workhere dealt with. The first riticism eals with the quantity f solution n whichtheculturesweregrowing. In nearly ll cases each culture onsisted f six plants growing ogetherin one culture ar containing 50 c.c. of solution. This means only42 c.c. of solutionperplant,and although he solutionswerechanged veryhreedays and although he plantswere mall, yetit is quite possible hat the plants were absorbing alts quite fast enough o alter appreciablyheproportion f the various alts in such a small quantity f solution.In the secondplace, no account s taken of the greatvariability f plants growing n watercultures nder xactly he same conditions. As only one culture ontaining ix plants was grownin the same solution t is verypossible that some of the differencesf dry weighton whichemphasis s laid, are within he imitsof theerror ue to this. But in the absence of any data inregard o probable rror t is not possible o say whether his s so.But regarded enerally t would appear that Mr Tottingham'swork has been most carefullycarried ut, and most of his conclusions re probablyustified. It is probably ifficult or hosewho have not conductedwork nvolving he use of water-cultureso realise the greatamountof abour nvolved n carrying o a successful onclusion uch a piece of work s that here con-sideredwiththe care that has evidently een bestowed pon t.To the ecologistof course he interest f this work ies very argely n the applicationof theresults btained o questions elating o the soil solution. How far t is permissible o argue fromeffectsroducedn water-cultureoproblems f the soil it is impossible t present o say. Thatsuch effectsmust have somebearingon soil problemss obvious,but we must wait beforewe cantelldefinitelyowhatextent ifferencesn concentrationf the soilsolutionorintheratiobetweendifferentubstances n the soil solutionare really operative n influencinghe distribution fvegetation.

WALTER STILES.

SOME RECENT WORK ON WATER AND MARSH VEGETATION(I) Snell, K. "Der Transpirationsstrom er Wasserpflanzen." Ber. d.deutsch. ot. Ges., 30, 1912, pp. 361-362.(II) Solereder, H. "Systematisch-anatomische ntersuchungendes Blattesder Hydrocharitaceen." Beih. z. Botan. Centralbl., 0, Abt. 1, 1913, pp. 24-104, 53 figures.

(III) Beyrer, H. "Beobachtungen iiber das Etiolment bei Wasserpflanzen."Jahresber. taats.-Gymn.n [etschen,14, 1913, pp. 3--16, 1 plate.(IV) MacDougal, D. T. " The determinativeaction of environic factorsupon Neobeckiaaquatica Green." Flora, 106, pp. 264-280, 14 text-figures.(V) Esenbeck, E. "Beitrage zur Biologie der GattungenPotaintogetonnidScirpus." Flora, 107, 1914, pp. 151-202, 59 text-figures.(VI) Gertz, A. "Pildammarnasvegetation fter orrlaggningen."Bot.Notiser,1913, pp. 113-130.The paperscitedabove are selected, ornoticehere,from hevery xtensive ecent iteratureon the ecologyof freshwaterquatic and marshvegetation, s a supplement o the collectivenoticeof papers on the subjectwhichappeared n an earliernumberof thisJOURNAL (1, 1913,pp. 107-115). This literatures growingo rapidly hat we have againomitted oticesof thosecontributions hich appear to be of minorgeneral nterest o the ecologist,nd have as a rule


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