About TERI The Bioresources and Biotechnology Division The...

About TERIA dynamic and flexible organization with a global vision and a local focus,TERI was established in 1974. While in the initial period the focus was mainlyon documentation and information dissemination activities, research activitiesin the fields of energy, environment, and sustainable development wereinitiated towards the end of 1982. The genesis of these activities lay in TERI’sfirm belief that efficient utilization of energy, sustainable use of naturalresources, large-scale adoption of renewable energy technologies, and reductionof all forms of waste would move the process of development towards the goalof sustainability.

The Bioresources and Biotechnology DivisionFocusing on ecological, environmental, and food security issues, the Division’sactivities include working with a wide variety of living organisms, sophisticatedgenetic engineering techniques, and, at the grassroots level, with villagecommunities. The Division functions through five areas—Centre for MycorrhizalResearch, Microbial Biotechnology, Plant Molecular Biology, Plant TissueCulture, and Forestry/Biodiversity. The Division is actively engaged inmycorrhizal research. The Mycorrhiza Network has specifically been created tohelp scientists across the globe in carrying out research on mycorrhiza.

The Mycorrhiza Network and the Centre for MycorrhizalCulture CollectionEstablished in April 1988 at TERI, New Delhi, the Mycorrhiza Network firstset up the MIC (Mycorrhiza Information Centre) in the same year, and theCMCC (Centre for Mycorrhizal Culture Collection) – a national germplasmbank of mycorrhizal fungi – in 1993. The general objectives of the MycorrhizaNetwork are to strengthen research, encourage participation, promoteinformation exchange, and publish the quarterly newsletter, Mycorrhiza News.

The MIC has been primarily responsible for establishing an informationnetwork, which facilitates information sharing among the network membersand makes the growing literature on mycorrhiza available to researchers.Comprehensive databases on Asian mycorrhizologists and mycorrhizal literature(RIZA) allow information retrieval and supply documents on request.

The main objectives of the CMCC are to procure strains of both ecto andVA mycorrhizal fungi from India and abroad; multiply and maintain these fungiin pure culture; screen, isolate, identify, multiply, and maintain nativemycorrhizal fungi; develop a database on cultures maintained, and providestarter cultures on request. Cultures are available on an exchange basis or onspecific requests at nominal costs for spore extraction/handling.

Vol. 13 No. 1April 2001

2 Mycorrhiza News 13(1) • April 2001

Mycorrhizal dependency, Part 1: selection of efficientmycorrhizal fungi

Sujan Singh ... 2

Research findingsMorphological and cytochemical study of the pelotonic and non-

pelotonic hyphae of the orchid Spathoglottis plicata ... 16Mass production of vesicular�arbuscular mycorrhizae using

different hosts ... 20A note on vesicular�arbuscular mycorrhiza development in

Albizia procera in root trainers ... 21

New approachesComparison of various techniques for recording AM colonization

... 22Molecular methods for determining intra- and interspecific

variations in ectomycorrhizal fungi ... 22

Centre for Mycorrhizal Culture CollectionMycorrhiza in sustainable agriculture ... 23

Recent references ... 25

Forthcoming events ... 28

Contents

* Compiled from TERI database – Riza

Mycorrhizal dependency, Part 1: selection of efficient mycorrhizal fungi

Sujan Singh*TERI, Darbari Seth Block, Habitat Place, Lodhi Road, New Delhi – 110 003, India

Mycorrhizal dependency of plants is defined vari-ously by mycorrhiza workers. Generally, when aparticular plant species/variety/cultivar/genotype inassociation with mycorrhiza produces greaterbiomass than that in the absence of mycorrhizaunder the same set of conditions and at any soilphosphorus level, then the species/variety/cultivar/genotype is considered as being dependent onmycorrhiza. To quantify the mycorrhizal depend-ency of a plant, it is necessary to conduct experi-ments in a soil that is most suitable for the growthof that plant and with a mycorrhizal fungus that ismost efficient on that plant. Also, it is necessary tofind the minimum soil solution phosphorus level atwhich maximum benefit may accrue to the plant, asgenerally the efficiency of mycorrhizal infection inthe plant is inversely proportional to the soil solu-tion phosphorus level. This write-up deals with theselection of efficient mycorrhizal fungi for differentplants.

Selection of efficient VA-mycorrhizalfungi for agricultural cropsCereal cropsRiceStudies conducted at the Department of Agricul-tural Microbiology, University of Agricultural Sci-ences, GKVK Campus, Bangalore, India, on rice(Cv ‘Prakash’) showed that Glomus intraradices wasmore efficient in improving grain yield than was G.fasciculatum. The increase in yield with G.intraradices was 11% and with G. fasciculatum was8% over uninoculated controls. With G. intraradicesinoculation, the amount of phosphate fertilizerusually applied to the rice crop may be reduced by50% (Secilia and Bagyaraj 1994).

Studies conducted at the Department ofBotany and Microbiology, Nagarjuna University,

Nagarjunanagar, Guntur, India, on eight varietiesof upland rice raised in pots with sterilized soilshowed that on all the varieties, Acaulospora spinosawas more effective in terms of levels of root coloni-zation, formation of vesicles/spores, plant biomass,and grain yield than was A. scrobiculata (Ammaniand Rao 1996).

WheatStudies conducted at the Biology Department,University of Western Sydney, Macarthur, Aus-tralia, on wheat var. swift showed that localaeroponically produced VAM inoculum of shearedroots of Sudan grass infected with trap culture ofGlomus intraradices was more effective in increasingyield at 0 kg phosphorus added per hectare underfield conditions. At 10 kg phosphorus per hectare,aeroponic VAM inoculum (both locally producedand imported from USA) was more efficient inincreasing the number of grains per spike than wasa single isolate sand culture of Gigaspora margarita,which was more efficient in increasing the numberof grains per spike at 20 kg phosphorus per hectarebut less effective than sheared-root inocula of G.intraradices in increasing 1000-grain weight at thislevel of phosphorus (Asif, Khan, Khan et al. 1995).

MaizeGreenhouse studies conducted at the Departmentof Biology, University of Ottawa, Canada, on maize(Zea mays) hybrid ‘Pioneer’ 3905 inoculated withGlomus aggregatum, G. etunicatum, G. mosseae, andG. versiforme showed that leaf mass and proteinconcentrations were higher and carbon percentagelower in plants inoculated with G. etunicatum incomparison with uninoculated controls and otherfungi. Nitrogen percentage was higher in plantscolonized by G. versiforme or G. aggregatum thanthose inoculated with G. mosseae. Sugar concentra-

Mycorrhiza News 13(1) • April 2001 3

tions in leaves were highest in plants inoculatedwith G. versiforme. However, the total shoot massand AM colonization (26%–72%) did not signifi-cantly differ among treatments (Boucher, Dalpe,and Charest 1999).

Studies conducted at the Department ofAgronomy and Soils, Clemson University,Clemson, USA, on corn (Zea mays) ‘Pioneer’3369A inoculated with five different AM fungi attwo pH levels in loamy sand soil showed that at pH6.1, using 6-month stored inoculum, shoot freshweights significantly increased with Gigasporagigantea and Glomus mosseae but not with G.macrocarpus var. macrocarpus, G. macrocarpus var.geosphorus or Gigaspora margarita. With 1-month-old inoculum, only G. mosseae showed significantfresh weight increase compared with the sterilecontrol. At pH 5.7, with 6-month-old inoculum,shoot fresh weights significantly increased with G.gigantea, G. mosseae, G. macrocarpus var. geosporus,and G. macrocarpus var. macrocarpus. With1-month-old inoculum, the fresh weight increasedwith G. gigantea only. All inoculated roots wereobserved to be over 70% infected for all VAM spe-cies (Struble, Skipper, and Smith 1979).

In experiments conducted at the Soil Microbi-ology Laboratory, G B Pant University of Agricul-ture and Technology, Pant Nagar, India,inoculation of maize with Gigaspora margarita,Glomus fasciculatum, and VAM-mixed endophyteindividually caused 54%, 50%, and 58% and rootcolonization respectively. Mycorrhizal root infec-tion decreased significantly with increasing levels ofP (Rathore and Singh 1995).

Pulse cropsGroundnutStudies conducted at the Texas AgriculturalExperiment Station, College Station, USA, on fivepeanut (Arachis hypogeae) cultivars grown at foursites and inoculated with four VAM fungi showedthat Glomus deserticola was the most effective spe-cies and G. mosseae was the least effective. Cultivar‘Florunner’ was the most responsive. An inoculummix of G. deserticola, G. intraradices, and G. mosseaeresulted in greater whole-plant weights than whensingle species were used. G. deserticola inoculationproduced shoot weights equivalent to those of themix (Taber, Smith, Smith et al. 1987).

Pot culture experiments conducted at the De-partment of Biosciences, Sri Sathya Sai Institute ofHigher Learning, Anantapur, Andhra Pradesh, India,on groundnut (Arachis hypogeae) showed that out ofeight indigenous VAM species tested, G. fasciculatumproduced the highest root–shoot length, dry weight,phosphorus content, and percent root colonization incomparison with other VAM species (Vijaykumar andBhirarvamurthy 1999).

Studies conducted at the Laboratory of PlantNutrition Faculty of Agriculture, Tokyo Universityof Agriculture and Technology, on cowpea, pigeon

pea and groundnut showed that these legumes re-sponded better to Glomus sp. than to Gigasporamargarita, in both topsoil (P = 72 ppm) and subsoil(P = 0.1 ppm). Despite low rates of root infection,shoot dry matter production was favoured bymycorrhizal inoculation and increases in shootbiomass were greater in subsoil than in topsoil. Ingroundnut, mycorrhization also raised P and Caconcentrations in topsoil (Ahiabor and Hirata1994).

Pigeon peaStudies conducted at the Tokyo University of Agri-culture and Technology on pigeon pea (Cajanuscajan) were similar to those for groundnut. How-ever, in pigeon pea, mycorrhization raised shootconcentrations of P and K in the topsoil (Ahiaborand Hirata 1994).

In studies conducted at the University ofGottingen, Grisebachstr, Germany, pigeon pea wasgrown in pots on non-sterilized phosphorus-fixingsoil supplied with phosphorus from three sources,namely single superphosphate (10, 30, 60 kg P/ha)rock phosphate with a total P of 10.7% (50, 150,300 kg/ha), and rock phosphate with a total P of12.1% (50, 150, 300 kg/ha). The soil containedtwo native species, Glomus albidum and G.intraradices. Four other VAM species originatingfrom the Cerrado ecosystem and G. manihotis werealso used. The inoculum was applied at the rate of2 g of air dry mixture of soil/roots/spores per plant.G. clarum proved to be the most effective fungusirrespective of the phosphorus source and level(Diederichs 1992).

SoybeanStudies conducted at the University of AgriculturalSciences, Dharwad, Karnataka, India, on soybeancv. AACS-58 sown in pots inoculated with Glomusfasciculatum or Gigaspora margarita and given 80,60, or 40 kg P

2O

5/ha showed that seed yield was

highest in plots inoculated with Gi. margarita + 80kg P2O5 (1413 kg/ha). G. fasciculatum + 80 kg P2O5/ha gave a seed yield of 1405 kg /ha. Soil inoculationwith mycorrhiza + 60 kg P

2O

5/ha gave a similar

seed yield as with no inoculation + 80 kg P2O5/ha(Lingaraju, Sreenivasa, and Babalad 1994).

Studies conducted at the Soil MicrobiologyResearch Group, Soil Science Division, Depart-ment of Agriculture, Bangkok, Thailand, onsoybean and mung bean grown in various fields ofThailand showed that of six VAM species tested,Acaulospora scrobiculata 22 was the most effectivefor soybean in growth and crop yield. Per cent rootcolonization, however, varied with time and phos-phorus nutrition in soybean (Vasuvat, Wadisirisak,Toomsen et al. 1987)

Mungbean (green gram)Studies conducted by the Department of Agricul-ture, Thailand, showed that of the six VAM fungitested, Glomus intraradices was the most efficient


for mung bean growth and crop yield. Unlike insoybean, per cent root colonization did not varywith time and phosphorus nutrition for mung bean(Vasuvat, Wadisirisali, Toomsen et al. 1987).

CowpeaStudies conducted at Tokyo University of Agricul-ture, Tokyo, showed that along with pigeon peaand groundnut, cowpea also responded better toGlomus spp. than Gigaspora margarita, both in top-soil and subsoil. Also, the effect of VAM on thiscrop was greater in subsoil than in topsoil as com-pared with the control (Ahiabor and Hirata 1994).

Studies conducted at the Agronomy Depart-ment, Clemson University, Clemson, USA, oncowpea grown on methylbromide-fumigated soilinoculated or not inoculated with Glomusetunicatum, G. claroideum, or native VAM fungi andalso with Bradyrhizobium strain 32 Hl showed thatinoculation with native VAM fungi gave higher topdry weight for 25-kg and 50-kg phosphorus levelsin the soil. Both G. etunicatum and G. claroideuminoculation gave a low top dry weight at lower Plevels and slightly higher top dry weights at high Plevels (Obura, Skipper, and Wagner 1987).

Black gram and green gramIn pot experiments conducted at the NagarjunaUniversity, Nagarjunanagar, Andhra Pradesh,India, black gram (Vigna mungo) and green gram(V. radiata) were grown in soil inoculated withAcaulospora spinosa, A. morrowae, and Glomusepigaeum (G. versiforme) and supplied with phos-phorus as superphosphate, rock phosphate, ortricalcium phosphate. The root colonization was68%–89% in black gram and 69%–93% in greengram. Colonization in both species was lowest inpots inoculated with A. morrowae + superphosphateand was best with G. epigaeum without addingphosphorus. Shoot dry weight, plant nitrogen andphosphorus concentrations, and 100-seed weightwere highest with super phosphate application + G.epigaeum inoculation (Rao and Rao 1996).

Vegetable cropsOnionA field trial was conducted at the Department ofHorticulture, College of Agriculture, Hyderabad,India, to study the response of onion to inoculationwith three VAM fungi at three phosphorus levelson a non-sterilized alfisol. Among the three VAMfungi, Gigaspora margarita proved to be the mosteffective inoculant in influencing the growth andperformance. Inoculation of onion in the nurserywith G. margarita at a phosphorus level of 17.5 kg/ha enhanced the number of leaves, dry matter ac-cumulation, uptake of phosphorus, and yield ofbulbs. Uptake of phosphorus and yield of onionbulbs with G. margarita inoculation at half the rec-ommended phosphorus level were significantlyhigher than that achieved by the uninoculated plant

at a full level of fertilization. G. calospora recordeda higher percentage of root infection at lower levelsof phosphorus (Ramana and Babu 1999).

CapsicumIn studies conducted at the Tata Energy ResearchInstitute, New Delhi, India, different types of nurs-ery inocula formulations, namely mixed indigenouscultures and Glomus intraradices, were comparedwith commercially available inocula in pot experi-ments using capsicum and Polianthus as testplants. Soil-based inocula and soil beads producedthe highest response in both crops. G. intraradicescaused the highest yield in capsicum (112% in-crease in fruit yield). Among the commercial in-ocula, only Mycorise enhanced fruit yield (11%) incapsicum. Sheared-root inocula of G. intraradicesresulted in high colonization (up to 68%), but theyield enhancement was lower than in the soil-basedformulations. The mixed indigenous inoculum pro-duced the highest number of spores and propaguleswhereas commercial inoculum produced the lowest(Gaur, Adholeya, and Mukerji 1998).

TomatoIn studies conducted at the Department of PlantPathology, University of Kentucky, Lexington,USA, tomato (cv. ‘Manpal’) seeds were inoculatedwith Glomus etunicatum or G. mosseae at severalinoculum levels (0.0, 0.1, 0.3, 0.5, 0.7, 1.0, or 10.0propagules per g of autoclaved fine sand soil). At2.5 weeks, significantly higher (P = 0.05) coloniza-tion index values (colonization per 0.5 g root freshweight sample × total root dry weight [g]) wereobtained for plants inoculated with G. mosseae thanfor those inoculated with G. etunicatum. A similarrelationship was observed at subsequent dates (at5, 6, 8 or 13 weeks). Conversely, G. etunicatuminoculation resulted in significantly higher (P =0.001) shoot dry weights and plant heights thandid G. mosseae inoculation or uninoculation plantsat 6 and 13 weeks. With cv. ‘Walter’, plants inocu-lated with G. mosseae had significantly greater shootdry weights and plant heights than those inoculatedwith G. etunicatum at 6 and 13 weeks (McGrawand Schenck 1981).

Fodder cropsAlfalfaIn studies conducted at the United States Depart-ment of Agriculture, Agricultural Research Service,Errc, Wyndmoor, USA, two cultivars of alfalfa(Medicago sativa), cv. ‘Giboa’ and cv. ‘Moapa’ 69,were inoculated in the glasshouse with three VAMfungi, namely Glomus mosseae, G. intraradices, andGigaspora margarita, to evaluate their relativeefficiencies. Along with alfalfa, Papsalum notatumwas also inoculated with these three fungi to de-scribe fungal parameters on a routine pot culturehost. Percentage root length of P. notatum colo-nized by VAM fungi increased from 10 to 21


weeks, and all fungi sporulated during this period.In alfalfa, only colonization by G. intraradices in-creased over that time period, and this was the onlyfungus to sporulate in association with alfalfa at 10weeks. G. mosseae did not sporulate after 16–21weeks despite having colonized 30%–35% of theroot length of both alfalfa cultivars. In vitro experi-ments in which RiTDNA-transformed roots ofalfalfa were inoculated with VAM fungi showednormal mycorrhizal formation by G. intraradicesand a hypersensitivity-like response to G. margarita.The colonized cells became necrotic and HPLCanalysis indicated increased concentrations ofphenolics and isoflavonoids in these root segments(Douds, Galvez, Becard et al. 1998).

CloverStudies conducted at the Soil Science and PlantNutrition Department, School of Agriculture, Uni-versity of Western Australia, Nedlands, Australia,on clover (Trifolium subterraneum) inoculated withthree species of Acaulospora in pots with 1.6 kgbrown sandy soil showed that A. trappei was moreeffective in increasing plant growth per unit of rootcolonization than was A. laevis and an unidentifiedAcaulospora sp. In the experiment, three harvests(3, 5, and 7 weeks after sowing) using six quantitiesof inoculum (10–250 g/pot) were obtained. For allthree fungi, increasing propagule number up tothree weeks increased per cent root colonization.This propagule effect decreased with time, and at 5and 7 weeks, there was no effect of propagulenumber on per cent root length colonized with A.trappei and the unidentified Acaulospora sp., butthere was marked difference with A. laevis at 5weeks, which disappeared at 7 weeks (Gazey,Abbott, and Robson 1992).

Finger milletStudies conducted at the GB Pant University ofAgriculture and Technology, Pant Nagar, UttarPradesh, India, on finger millet (Eleusine coracana)showed that out of six different VAM fungi(Glomus caledonicum, G. mosseae, G. fasciculatum,G. epigaeum, G. versiforme, Gigaspora calsospora, andGi. margarita) tested, maximum root colonizationand mycorrhizal efficiency were observed in plantsinoculated with G. caledonicum (Tewari, John, andTandon 1993).

GuarPot experiments conducted at the Department ofBotany, JNV University, Jodhpur, Rajasthan, India,on guar (Cyamopsis tetragonoloba) showed that outof six VAM fungi (Acaulospora morrowae, Glomusconstrictum, G. fasciculatum, G. mosseae, Sclerocystisrubiformis, and Scutellospora calospora), all collectedfrom the rhizosphere of guar, G. fasciculatumcaused a twofold increase in protein contents and a50% increase in total sugar contents in pods ascompared with controls (Mathur and Vyas 1995).

Tuber cropsAmorphophallus, Cassava, and ColocasiaIn studies conducted at the Centre for AdvancedStudies in Botany, University of Madras, TamilNadu, India, cassava (Manihot esculenta),Amorphophallus paeonifolius, and kachalu (Colocasiaesculenta) were grown in both pots and field andwere inoculated with Gigaspora albida, Pisolithustinctorius, Glomus mosseae, or G. aggregatum. Jointinoculations with G. mosseae and G. aggregatumgave the highest tuber yield per plant in all threecrops (Ganesan and Mahadevan 1994).

Studies conducted at the College of Agricul-ture, Velleyani, Trivandrum, Kerala, India, on cas-sava in pot cultures showed that inoculation withGlomus fasciculatum gave the highest shoot and rootdry weight, plant height, and VAM infection whencompared with those inoculated with G. mosseae,G. constrictum, G. etunicatum, Acaulosporamorroweae, and controls (Sivaprasad, Sulochana,and Nair 1990).

Fruit cropsPineapple, papaya, and bananaStudies conducted at the Departmento deProteccion Vegetal, Centre de Investigation YTechnologia Agrarias La Laguna, Canary Islands,Spain, on papaya (Carica papaya), pineapple(Ananas comosus), and banana (Musa acuminata)showed that all the species were highly responsiveto mycorrhizal conditions under greenhouse as wellas field conditions when the AM inoculum wasGlomus endophytes. Glomus fasiculatum was themost effective in improving plantlet growth andnutrition. Acaulospora sp. was ineffective andScutellospora sp. was effective with banana only(Jaizme-Vega and Azcon 1995).

StrawberryIn studies conducted at the SLVA Ahrweiler,Walporzheimer Str. 48, Bad Neunahr-Ahrweiler,Germany, micropropagated and runner-propagatedstrawberry (Fragaria × ananassa) cv. Elvira andElsanta were inoculated with Glomus etunicatum, Gversiforme (two isolates), G. mosseae, G. versiforme33-366 increased yield and runner production de-pending on cultivar and propagation method.Micropropagated plants had higher mycorrhizationrates and suffered less from infection byPhytophthora cactorum than runner-propagatedplants (Taube-Baab and Baltruschat 1996).

Other plantsRajnigandhaIn studies conducted at the Tata Energy ResearchInstitute, New Delhi, India, on evaluation of differ-ent nursery inocula formulations, soil-based in-ocula and soil beads produced the highest responsein both rajnigandha (Polianthus tuberosa) and capsi-cum. Glomus intraradices gave the highest increase


in spike length (45%) in rajnigandha. Among thecommercial inocula, only Mycorise enhanced spikelength (33%). Overall, AM colonization was higherin rajnigandha than in capsicum. Sheared-rootinoculum of G. intraradices resulted in high rootcolonization (up to 68%), but the yield enhance-ment was lower than with soil-based formulation.The mixed indigenous inoculum produced thehighest number of spores and propagules whereasthe commercial inoculum produced the lowest(Gaur, Adholeya, and Mukerji 1998).

PassifloraGreenhouse trials conducted at the UniversidadNacional de Colombia, Sede Palmira, Colombia(1200 m above sea level, at 24 ºC) on Passifloraligularis plants grown in soil sterilized and inocu-lated or not inoculated with Acaulospora foveata, A.longula, or Glomus occultum at pH 4.8 showed thatA. longula was the most effective VAM fungus(Rodriguez, Hurtado, and Sanchez de Prager1995).

Selection of efficient mycorrhizal fungifor tree cropsSelection of VA mycorrhizal fungiVA mycorrhizal fungi form symbiotic associationsmainly with hardwood trees of both forest and hor-ticultural species.

Horticulture treesAppleStudies conducted at the Department of Horticul-ture, Ohio State University, Columbus, Ohio,USA, on apple (Malus × domestica) showed thatGlomus mosseae exceeded G. macrocarpum in therate of initial colonization, amount of maximumcolonization, and persistence of arbuscules andexternal hyphae. Colonization by each fungus re-duced fresh weight 11 days after inoculation butincreased shoot and total fresh weight by day 38.Relative height growth rate and colonization atharvest of plants inoculated with G. macrocarpumwere less than those of plants inoculated with G.mosseae or G. mosseae + G. macrocarpum, butgreater than controls (Miller, Bodmer, andSchuepp 1989).

CitrusIn studies conducted at the United States Horticul-tural Research Laboratory, United States Depart-ment of Agriculture, Orlando, USA, rough lemon(Citrus jambhiri) was grown on fine sand amendedwith individual and combined populations ofGlomus mosseae, G. etunicatum and G. fasciculatum(900 chlamydospores for a 15-cm pot). Mean plantheights for G. mosseae, G. etunicatum, and G.fasciculatum were 21.6, 34.6, and 52.5 cm, topweights were 6.4, 14.2, and 25.2 g, root weightswere 14.3, 32.4, and 63.4 g, and chlamydosporeswere 67, 4, and 73/25 g soil respectively. All

growth parameters were significantly different (P =0.01) from each other and from uninoculated con-trols. In all combinations of two species, heightvaried between 38 cm and 42 cm, top weight be-tween 18 g and 20 g, and root weight between 21 gand 26 g. None of the individual growth valuesdiffered significantly within each parameter. Sporu-lation in tests with G. mosseae and G. etunicatumwas 5 and 0 chlamydospores per 25 g soil, with G.mosseae and G. fasciculatum was 6 and 25chlamydospores per 25 g soil, and with G.etunicatum and G. fasciculatum was 5 and 6chlamydospores per 25 g soil respectively. In thetest with all the three species, height was 33.4 cm,top weight 13.6 g and root weight 15.7 g. In thistest, only G. mosseae sporulated and that too onlywith one spore per 25 g soil (Nemec 1979).

Studies conducted at the Department of Agri-culture Microbiology University of AgriculturalSciences, GKVK, Banglore, India, on the responseof Trifoliate orange (Poncirus trifoliata) to 18 VAMfungi showed that Glomus velum, G. caledonicum, G.marredum, G. macrocarpum and Acaulospora sp.were the most effective in improving growth andnutrition, resulting in greater plant height, stemdiameter, plant biomass, P, Zn, and Cu context.All the five fungi selected were statistically equiva-lent in improving growth and nutrition of P.trifoliate. Inoculation of the root stock with thesefungi produced plants ready for budding in 14–15months as compared with 22–24 months withoutmycorrhizal inoculation.

ZizyphusStudies conducted at the Department of Botany,JNU University, Jodhpur, Rajasthan, India, onZizyphus mauritiana raised in pots and inoculatedwith Glomus fasiculatum, G. mosscae, or Scutellosporacalospora showed that G. fasciculatum was signifi-cantly more efficient than the other two VAM fungiin increasing seedling biomass and uptake of nitro-gen, phosphorus, potassium, calcium, and magne-sium after 98 days of inoculation. All fungi werecollected from the rhizosphere of Z. mauritiana andthe inoculum used in the experiment was in theform of roots of Cenchrus ciliaris colonized by eachVAM fungus (Mathur and Vyas 1996).

Forest treesTeakStudies conducted at the University of AgriculturalScience, GKVK Campus, Bangalore, India, on teak(Tectona grandis) stumps planted in polybagsshowed that on the basis of various plant growthparameters and nutritional status of the plants,Glomus leptotichum was the best AM symbiont forteak (Rajan, Reddy, and Bhagyaraj 2000).

RubberStudies conducted at the Rubber Research Insti-tute of Malaysia, Kuala Lumpur, Malaysia, on rub-ber (Hevea brasilienseis) (PB5/51) seedling root


stock showed that after 22 weeks of growth in steri-lized soil, with phosphorus added or not added,only 2 of the 10 VAM fungi (Glomus clarum,Glomus sp.) showed a significant increase in shootdry weight compared with uninoculated controls,and none showed an increase in root fresh weight.Stem diameter and height growth from inoculationwere not affected by any of the endophytes butinfection by Acaulospora dilatata, A spinosa,Gigaspora margarita, Entrophospora colombiana, andan unknown Glomus sp. increased foliar phospho-rus concentrations. The amount of root infectedwas not related to improved plant growth (Ikram,Mahmud, and Othman 1993).

NeemStudies conducted at the Department of Agricul-tural Microbiology, GKVK, Bangalore, India, onneem (Azadirachta indica) seedlings inoculated at45 days with nine VAM fungi showed that neemseedlings responded best to inoculation withGlomus mosseae followed by G. fasciculatum as re-gards plant height, stem girth, biomass, phospho-rus content, zinc concentration, biovolume index,mycorrhizal root colonization, and spore number inthe root-zone soil (Sumana and Bagyaraj 1999).

BabulStudies conducted at the Department ofAgroforestry, Haryana Agricultural University,Hissar, Haryana, India, on babul (Acacia nilotica)seedlings inoculated with Glomus constrictum, G.mosseae, and G. gilmorei (collected from old babulplantations) singly or in combination in a phospho-rus-deficient soil showed that G. mosseae combinedwith G. gilmorei was the most effective in improvingcollar diameter, root–shoot dry weight, and lengthin comparison with uninoculated plants. G.constrictum was the least effective fungus (Mandaland Kaushik 1994).

Selection of ectomycorrhizal fungi �hardwoodsOakIn studies conducted at the Department of ForestResources, College of Forestry, University of Min-nesota, USA, seedlings of three species of oak weregrown in containers (receiving 75 ml Hoagland’ssolution per seedling) and inoculated with five iso-lates of Pisolithus tinctorius, three of Suillusgranulatus, and one each of S. luteus, Thelephoraterrestris, and Cenococcum geophilum. Root coloniza-tion differed significantly among fungal isolates andamong oak species. T. terrestris was most effectivein increasing nutrient uptake by the three oaks col-lectively. Individually, S. granulatus 263 was themost effective with Quercus velutena, S. luteus wasthe most effective with Q. alba, and C. geophilumwas the most effective with Q. robur (Mitchell,Cox, Dixon et al. 1984).

EucalyptusIn studies conducted at the CSIRO, Division ofForestry, Wembley, Australia, Eucalyptus globulusand E. diversicolor were grown in yellow sand at twolevels of phosphorus (4 and 12 mg phosphorus/ kgsand). Both the tree species were inoculated with16 ectomycorrhizal fungi, which included Paxillusmuelleri, Cortinarius globuliformis, Thaxterogaster sp.;Hysterangum inflatum, Hydrangium carneum,Hymenogaster viscidus, H. zeylanicus, Setchelliogastersp.; Laccaria laccata, Scleroderma verrucosum,Amanita xanthocephala, Descolea maculata, andPisolithus tinctorus. At the time of harvest (100days), uninoculated seedlings and those inoculatedwith P. muelleri and C. globuliformis had a low levelof contamination from an unknown mycorrhizalfungus. Thaxterogaster sp. and H. inflatum devel-oped a superficial type of mycorrhiza. All othersformed typical pyramidal ectomycorrhizas. The dryweights of inoculated and uninoculated E. globulusseedlings at 12 mg phosphorus/kg soil did not differwhereas several isolates caused growth depressionin E. diversiecolor. At 4 mg phosphorus, growthincreases in inoculated seedlings were 0–13 timeshigher than the growth rate of uninoculated seed-lings. P. tinctorius produced the largest growth in-creases in both Eucalyptus species. In general,isolates that developed more extensive mycorrhizason roots produced the largest growth responses toinoculation. Isolates that increased plant growthalso increased phosphorus uptake by the plant.Seedlings inoculated with L. laccata and S.verrucosum retained more phosphorus in their rootsthan did plants associated with the other fungalisolates (Burgess, Malajczuk, and Grove 1993).

Selection of ectomycorrhizal fungi �conifersPinesStudies conducted at the Institute National delaRecherche Agronomique, Centre de RechercheForestiere, Champenoux, Seichamps, France, onPinus sylvestris at different levels of phosphorusshowed that even a low intensity of infection byLaccaria laccata greatly stimulated P. sylvestrisgrowth in comparison with Thelephora terrestris andHebeloma crustuliniforme. The ability of L. laccata toincrease P. sylvestris growth seemed to be morerelated to its capacity to produce growth sub-stances than to its capacity to stimulate phosphorusuptake. The poor efficiency of H. crustuliniforme incomparison with L. laccata at any level of phospho-rus could result from differences in diversion ofcarbohydrates from the host to fungal structures(Tyminska, Tacon, and Chadoeuf 1986).Studies conducted at the Centre Advanced Studies inBotany, University of Madras, India, on Pinus patulaseedlings showed that up to the first six months,seedlings inoculated with Thelephora terrestris showedbetter growth, mycorrhizal development, and dry


matter production, but at later stage, seedlingsinoculated with Laccaria laccata grew better of thetwo fungi. Under experimental conditions, L.laccata was found better for P. patula nurseries(Natarajan and Reddy 1994).

Studies conducted at the University of Murcia,Faculty of Biology Vegetal Botany, Murcia, Spain,on Pinus halepensis inoculated with three differentbasidiospore concentrations of Pisolithus arhizus,Rhizopogon roseolus, and Suillus collinitus showedthat highest mean values of height, dry weight, andpercentages of ectomycorrhizas were obtained withseedlings inoculated with P. arhizus in a sterilesubstrate (Torres and Honrubia 1994).

Douglas firIn studies conducted at the Department dePatologia Vegetal, Institute Recerca i TechnologiaAgroaliamentarie Cabrils, Spain, 36 isolates from27 species of native ectomycorrhizal species weretested with Douglas fir (Pseudotsuga menziesii) and13 species were tested with two native and fourintroduced species of conifers in pure culture syn-thesis. Only eight fungi colonized over 50% of theshort roots. Laccaria laccata, Lyophyllum decastes,Pisolithus tinctorius, and Scleroderma citrinum formedabundant mycorrhizas in all conifers tested.Twenty-three fungal isolates from 18 speciesformed ectomycorrhiza with Douglas fir. Ninefungi did not form ectomycorrhiza even thoughsome of them were collected in pure stands ofDouglas fir. Lactarius deliciosus, Rhizopogon sp., andSuillus luteus showed the highest host specificity(Parlade, Alvarez, and Pera 1996).

Sitka spruceIn studies conducted at the United States Depart-ment of Agriculture, Forest Service, Pacific North-west Forest Range Experiment Station, Portland,Oregon, USA, containerized sitka spruce (Piceasitkensis) seedlings were inoculated at sowing withinoculum of Pisolithus tinctorius, Laccaria laccata,Astraeus pteridis, Amanita pantherina or Cenococcumgeophilum, all common in sitka spruce forests. Theinoculum was prepared in vermiculite culture andcombined with a peat vermiculite potting mixturein a 1:7 ratio. The seedlings were grown for sixmonths in 66-cm3 cells in a greenhouse atCorvallis, Oregon (Location 1), environmentalgrowth chambers at Juneau, Alaska (Location 2),and a small production nursery greenhouse atPetersburg, Alaska (Location 3). At locations 1, 2and 3, respectively 100%, 100%, and 85% seed-lings inoculated with L. laccata formed mycorrhizaand 100%, 98%, and 80% inoculated with C.geophilum formed mycorrhizae. Mycorrhizal shortroots at locations 1, 2, and 3 respectively were92%, 77%, and 38% for L. laccata and 91%, 12%,and 7% for C. geophilum. Positive mycorrhiza for-mation was not detected at any location with the

other test fungi. Analysis of variance for growthparameters within each location for seedlings colo-nized by L. laccata and C. geophilum showed signifi-cant differences ( P = 0.05) only for root collardiameter, root weight, and total weight at location1. Tukey tests for differences among treatmentmeans for these variables were not significant ( P =0.05); however, for all three variables, mean valuesfor the control seedlings were greater than those forseedlings colonized by either fungus (Shaw andMolina 1979).

Two sets of experiments were conducted at theDepartment of Botany, University of Guelph,Guelph, Ontario, Canada, on Picea mariana, seedsof which were collected from an upland site withsandy soil and lowland site with peaty soil for rais-ing seedlings. In one set grown under aseptic con-ditions in test tubes, seedlings were inoculated withHebeloma cylindrosporum, Laccarier laccata andPaxillus involutus; in the other set, the seedlingswere inoculated with Laccaria bicolor, L. laccata,L. proxima or uninoculated agar plugs. Differencesbetween fungal treatments were pronounced. BothL. bicolor and L. laccata initiated good ectomycorrhizaformation and increased seedlings growth(Thomson, Mathes-Sears, and Peterson 1990).

TamarackStudies conducted at the Department of ForestScience, University of Alberter, Edmonton, Canadaon containerized seedlings of tamarack (Larixlaricina) (representing four provenances and 17open-pollinated families) inoculated with fourectomycorrhizal fungi showed that during an 18-week period, Laccaria laccata, Cenococcumgeophilum, and Pisolithus tinctoruis formedectomycorrhizae with 60%, 7%, and 12% of thetotal short roots respectively. Suillus granulatusfailed to form any mycorrhiza. Overall, the bestgrowth was found in seedlings inoculated L. laccata(Zhu and Nauratil 1987).

FernIn studies conducted at the Centre de Rechercheen Biologie Forestiere, Universite Laval, Quebec,Canada, rooted plantlets of in vitromicropropagated fern, Nephrolepis exaltata var.Whitmanii were transferred to pots containing abrown peat-based mix and simultaneously inocu-lated with Glomus intraradices, G. clarum, G.vesiculiferum, or G. versiforme. Root colonization byG. intraradices and G. clarum was more rapid andextensive as compared with that of the other twofungi. The high-phosphorus control performedbetter than other treatments except for the fondnumber, which did not differ significantly. Amongmycorrhizal treatments, plants inoculated with G.vesiculiferum showed the most significant increasein growth as compared with low-phosphorus con-trol (Ponton, Piche, Parent et al.1990).

Mycorrhiza N

ews 13(1)

•A

pril 2001

9

Plant species

Agriculture cropsAllium cepa (onion)

Amorphophallus paeonüfolius

Ananas comosus

Arachis hypogeae

Acacia nilotica

Acaciaauriculaeformis

Acer saccharum(sugar maple)

Azadirachta indica(neem)

Cajanus cajan

Capsicum

Carica papaya

Colocasia esculenta

Efficient mycorrhizal fungus/ fungi

Gigaspora margarita

G. calospora

Glomus mosseae +G. aggregatum

Glomus fasciculatum

Glomus sp.

Glomus deserticola

Glomus fasciculatum

Glomous mosseae + Gigasporagilmorei

Glomus etunicatum, G. macrocarpum

Glomus macrocarpum

Glomus mosseae

Glomus clarum

Glomus etunicatum

Glomus intraradices

Glomus fasciculatum

Glomus mosseae +G. aggregatum

Mycorrhizal fungi tested

G. margarita, G. calospora, and a known VAMfungus

G. margarita, G. calospora, and a known AMfungus

G. mosseae, G. aggregatum, Gigaspora albida,Pisolithus tinctorius

G. fasciculatum, Glomus sp., Acaulospora sp.,Scutellospora sp.

G. etunicatum , Gigaspora margarita

G. deserticola, G. mosseae, Glomusintraradices, and another VAM fungus

G. fasciculatum and seven other VAM fungi

G. mosseae, G. constrictum, Gigasporagilmorei, and their combinations

G. etunicatum, G. macrocarpum,G. fasciculatum, G. mosseae

G. macrocarpum, G. clarum, G. epigaeum,G. mosseae

Glomus mosseae, G. fasciculatum, and sevenother VAM fungi

G. fasciculatum and seven other VAM fungi

G. etunicatum , Gigaspora margarita

G. intraradices, indigenous mixed culture,commercial inoculum (Mycorise & C)

G. fasciculatum, Glomus sp., Acaulospora sp.,Scutellospora sp.

G. mosseae, G. aggregatum, Gigaspora albida,Pisolithus tinctorius

Parameters evaluated

Leaf number, dry matter, P-uptake, bulbyield

Root colonization

Tuber yield/plant

Plant growth, nutrition

Shoot dry matter, nutrient uptake

Plant growth

Root�shoot length, dry weight, P-content,root colonization

Collar diameter, root�shoot dry weightand length

Height, diameter, biomass, P-uptake

Leaf number, mean lateral root number

Plant height, stem girth, biomass, P-con-tent, zinc concentration, root colonizations

Plant growth

Shoot dry matter, nutrient uptake

Fruit yield

Plant growth, nutrition

Tuber yield/plant

References

Ramana and Babu (1999)

Ramana and Babu (1999)

Ganesan and Mahadevan(1994)

Jaizme-Vega and Azcon (1995)

Ahiabor and Hirata (1994)

Taber, Smith, Smith et al.(1987)

Vijay Kumar and Bhiravamurthy(1999)

Mandal and Kaushik (1994)

Aggangan, Lorilla, and Cruz(1990)

Reid, Parker, Mitchell et al.(1988)

Sumana and Bagyaraj (1999)

Diederichs (1992)

Ahiabor and Hirata (1994)

Gaur, Adholeya, and Mukerji(1998)

Jaizme-Vega and Azcon (1995)

Ganesan and Mahadevan(1994)

(Continued)

Appendix 1 List of efficient VA mycorrhizal fungi for different agricultural species


Plantspecies

Cucumissativus

Cyam

opsistetragonoloba

Eleusine

coracana

Fragariaxananassa

(microprop

agated)

Glycine

max

Lycopersicum

esculentum

cv.M

anapal

cv.W

alter

Manihot

esculenta

(cassava)

Manihot

esculenta

(cassava)

Medicagosativa

Musaacum

inata

Oryza

sativa

cv.P

rakash

Orzya

sativa

(uplandrice)

Passifloraligularis

Persea

americana

Phaseolusmungo

(mung

bean)(Vigna

radiata)

Polianthestuberosa

(rajnigand

ha)

Efficient

mycorrhizalfungus/fungi

Glomus

caledonium

Glomus

fasciculatum

Glomus

caledonicum

Glomus

versiform

e36

-366

Acaulspora

scrobiculata

Glomus

etunicatum

G.m

osseae

G.m

osseae

Glomus

fasciculatum

G.m

osseae

+G.aggregatum

Glomus

intraradices

Glomus

fasciculatum

Glomus

intraradices

Acaulosporaspinosa

Acaulosporalongula

Glomus

fasciculatum

Glomus

intraradices

Glomus

intraradices

Mycorrhizalfungitested

G.caledonium,G

lomus

sp.

G.fasciculatum,G

.mosseae,A

caulospora

morrowae,G

.constrictum,S

clerocystis

rubiform

e,Scutellosporacalospora

G.caledonicum

,G.m

osseae,G

.fasciculatum,

G.epigaeum(G.versiform

e),G

igaspora

calospora,

G.m

argarita

G.versiform

e(2

isolates),G.m

osseae,

G.e

tunicatum

A.scrobiculata,G

lomus

intraradices,a

ndfour

otherV

AMfungi

Glomus

etunicatum

,G.m

osseae

Glomus

etunicatum

,G.m

osseae

Glomus

etunicatum

,G.m

osseae

G.fasciculatum,G

.mosseae,G

.constrictum,

G.etunicatum,A

caulospora

morrowea

G.m

osseae,G

.aggregatum,G

igaspora

albida,

Pisolithustin

ctorius

G.intraradices,G.m

osseae,G

igaspora

margarita

G.fasciculatum,G

lomus

sp.,Aculospora

sp.,

Scutellosporasp.

G.intraradices,G.fasciculatum

Acaulosporaspinosa,

A.scrobiculata

A.longula,

A.foveata,

Glomus

occultu

m,n

ative

soilVA

Mfungi

G.fasciculatum,G

lomus

sp.,Acaulosporasp.,

Scutellosporasp.

G.intraradices,Acaulosporascrobiculata,a

ndfour

otherfungi

G.intraradices,indigenous

mixed

cultu

res,

commercialinoculum

(Mycorise)

Parametersevaluated

P-up

take

Proteinandsugarcontentsinpods

Mycorrhizalefficacy,root

colonizatio

n

Yield,

productio

nofrunners

Plantgrowth,cropyield

Shootd

ryweight,planth

eight

Root

colonizatio

n

Shootw

eight,planth

eight

Root

colonizatio

n,plantw

eight,shoota

ndroot

dryweight

Tuberyield/plant

Root

colonizatio

n,fungalsporulation

Plantgrowth,n

utrition

Grainyield

Plantbiomass,grainyield,rootcolonization

Root

diam

eter,rootw

eight,dryweighto

faeria

lparts,totalroot

length,spore

numberinsoil

Plantgrowth,n

utrition

Plantgrowth

,cropyield

Fruityield,spike

length

References

Jonera

ndJokobsen

(199

4)

Mathura

ndVyas

(199

0)

Tewari,Johri,andTandon

(199

3)

Taube-Ba

abandBa

ltruschat

(199

6)

Vasuvat,Nopam

ornbodi,and

Tham

surakul(19

87)

McG

rawandSchenck(198

1)

McG

rawandSchenck(198

1)

McG

rawandSchenck(198

1)

Sivaprasad,S

ulochana,and

Nair

(199

0)

Ganesan

andMahad

evan

(199

4)

Douds,G

alvez,Be

card

etal.

(199

8)

Jaisme-Vega

andAzcon(199

5)

Secilia

andBa

gyaraj(199

4)

Ammaniand

Rao(199

6)

Rodriguez,Hurtado,a

ndSa

nchez

(199

5)

Jaizme-Vega

andAzcon(199

5)

Vasuvat,Wadisirisak,Toom

sen

etal.(19

87)

Gaur,Ad

holeya,a

ndMukerji

(199

8)

Appendix1

Continued


Trifolium(clover)

Triticumaestivum

wheat

(var.swift)

Vignamungo

(black

gram

)

Vignaradiata(green

gram

)

Vignaunguiculata

Vignaunguiculata

cv.K

atum

anuK8

0

Vignaunguiculatacv.

CaliforniaNo.5blackeye

Zeamays(Pioneer

3905

)

Nephrolepisexaltata

var.

whitm

anii(fern)

micropropagated

plants

Tree

crops

Malus

×domestica(app

le)

Hevea

brasiliensis

Leucaena

leucocephala

(K-8,K

-28)

Tectonagrandis

Zizyph

usmauritiana

Citrus

jambhiri

(rough

lemon)

Persea

american

(avocado)

Acaulosporatrappei

Glomus

intraradices

Glomus

epigaeum

(G.versiform

e)

Glomus

epigaeum

(G.versiform

e)

G.etunicatum

G.e

tunicatum

G.c

laroideum

Glomus

etunicatum

Glomus

intraradices,G

.clarum

Glomus

mosseae

Glomus

clarum

,Glomus

sp.

Glomus

mosseae

Glomousleptotichus

Glomus

fasciculatum

Glomus

fasciculatum

Glomus

fasciculatum

A.trappei,A.

laevis,A

caulospora

sp.

G.intraradices,Gigaspora

margarita

G.epigaeum,A

caulospora

spinosa,

A.morrowae

G.epigaeum,A

caulospora

spinosa,

A.morrowae

G.etunicatum,G

igaspora

margarita

Glomus

etunicatum

,G.claroideum,G

lomus

sp.

Glomus

etunicatum

,G.claroideum,G

lomus

sp.

G.etunicatum,G

.mosseae,G

.aggregatum,

G.versiform

e

G.intraradices,G.clarum,G

.versiculiferum

,G.versiform

e

G.m

osseae,G

.macrocarpum

G.clarum,G

lomus

sp.,

A.spinosa,Gigaspora

margarita,

Entrophosporacolombiana,

Glomus

sp.,and

threeotherspecies

G.m

osseae

andsixotherV

AMfungi

Not

mentio

ned

G.fasciculatum,G

.mosseae,S

cutellospora

calospora

G.fasculatum,G

.mosseae,G

.etunicatum

Glomus

fasciculatum

,Acaulospora

sp.,

Scutellosporasp.

Root

colonizatio

n

Plantyield,n

umbero

fgrains/

spike

Root

colonizatio

n,shootd

ryweight,Nand

Pconcentrations

Root

colonizatio

n,shootd

ryweight,Nand

Pconcentrations

Shootd

rymatter,nutrient

uptake

Topdryweight

Topdryweight

Leafmass,proteinconcentrations

Root

infection

Root

colonizatio

n,planth

eight,plant

grow

th

Dryweight

Seedlinggrow

th

Seedlinggrow

thparameters

Growth,rootcolonization,

nutrient

uptake

Planth

eight,topweight,root

weight,

chlamydosporesinsoil

Plantletgrowth

andnutrition

Gazey,A

bbott,andRo

bson

(199

2)

Asif,

Khan,K

hanetal.(19

95)

RaoandRa

o(199

6)

RaoandRa

o(199

6)

Ahiabora

ndHira

ta(199

4)

Obura,S

kipp

er,a

ndWagner

(198

7)

Obura,S

kipp

er,a

ndWagner

(198

7)

Boucher,Dalpe,and

Charest

(199

9)

Ponton,P

iche,P

arenteta

l.(199

0)

Miller,B

odmer,a

ndSchuepp

(198

9)

Ikram,M

ahmud,and

Othman

(199

3)

Byra

Redd

yandBa

gyaraj(198

8)

Rajan,

Redd

y,andBa

gyaraj

(200

0)

Mathura

ndVyas

(199

6)

Nem

ec(197

9)

Jaizme-Vega

andAzcon(199

5)

12

Mycorrhiza N

ews 13(1)

•A

pril 2001

Plant species

Eucalyptus globulus

Eucalyptus diversicolor

Larix laricina(four provenances)

Picea engelemanü(Engleman spruce)

Picea sitkensis (Sitka spruce)

Pinus halepensis

Pinus patula (6 months)

Pinus patula (after 6 months)

Pinus sylvestris

Pinus sylvestris

Pinus contorta (Lodgepolepine)

Efficient mycorrhizal fungus/ fungi

Pisolithus tinctorius

Pisolithus tinctorius

Laccaria laceata

Amphinema bysoides and E. strain

Laccaria laccata, Cenococcumgeophilum

Pisolithus arhizus

Thelephora terrestrus

Laccaria laccata

Laccaria laccata

Amanita muscaria, Lactarius rufus

Suillus sp.

Mycorrhizal fungi tested

P. tinctorius, Paxillus muelleri, Cortinariusglobuliformis, Thexterogaster sp., Hysterangiuminflatum, Hydnangium carneum, Hymenogasterviscides, H. zealanicus, Setchelliogaster sp.,Laccaria laccata, Scleroderma verrucosum,Amanita xanthocephala, Descolea maculata,and three other fungi

P. tictorius, Paxillus muelleri, Cortinariusglobuliformis, Thexterogaster sp., Hysterangiuminflatum, Hydnangium carneum, Hymenogasterviscides, H. zealanicus, Setchelliogaster sp.,Laccaria laccata, Scleroderma verrucosum,Amanita xanthocephala, Descolea maculata,and three other fungi

L. laceata, Cenococcum geophilum, Pisolithustinctorius, Suillus granulatus

A. bysoides, E. strain, Thelephora terrestris,Cenococcum geophilum, Mycelium radices-atro-viren, Tomentella sp., Laccaria spp.,Hebeloma, Tuber, Suillus, Rhizopogon

L. laccata, C. geophilum, Pisolithus tinctorius,Astraeus pteridis, Amanita pantherina

P. arhizus, Rhizopogon rosovlus, Suilluscollinitus

T. terrestris, Laccaria laccata

T. terrestris, Laccaria laccata

L. laccata, Hebeloma crustuliniforme,Thelephora terrestris

A. muscaria, L. rufus, Laccaria laccata,Hebeloma crustuliniforme

E. strain, Amphinema bysoides, Thelephoraterrestris, Cenococcum geophilum, Myceliumradices atrovirins, Tomentella spp., Laccariaspp., Hebeloma, Tuber suillus, Rhizopogon

Parameters evaluated

Plant growth, P-uptake

Plant growth, P-uptake

Overall growth

Diameter growth

Mycorrhizal infection

Plant height, dry weight, mycorrhizalinfection

Growth, dry matter mycorrhizal infection

Growth, dry matter mycorrhizal infection

Plant growth

Seedling growth

Diameter growth

References

Burgess, Malajczuk, and Grove(1993)

Burgess, Malajczuk, and Grove(1993)

Zhu and Navratel (1987)

Hunt (1990)

Shaw and Molina (1979)

Torres and Honrubia (1994)

Natarajan and Reddy (1994)

Natarajan and Reddy (1994)

Tyminska, Tacon, and Chadoeuf(1986)

Stenstrom (1990)

Hunt (1990)

Appendix 2 List of efficient ectomycorrhizal fungi for tree crops


Pseudotsugamenzcesu

(Douglas

fir)

Quercus

alba

Q.rubor

Q.velutina

Quercus

petracea

Q.rubra

Rhizopogon

spp.

Thelephora

terrestris

Thelephora

terrestris

Thelephora

terrestris

Paxillusinvolutus

Paxillusinvolutus

E.strain,A

mphinem

abysoides,Thelephora

terrestris,C

enococcumgeophilum,M

ycelium

radicesatrovirin

s,Tomentella

spp.,Laccaria

spp.,H

ebelom

a,Tubersuillus,Rh

izopogon

Pisolithustin

ctorius(5

isolates),Su

illus

gram

ulatus

(3isolates),Su

llusluteus,

Thelephora

terristris,C

enococcumgeophulum

Pisolithustin

ctorius(5

isolates),Su

illus

gram

ulatus

(3isolates),Su

llusluteus,

Thelephora

terristris,C

enococcumgeophulum

Pisolithustin

ctorius(5

isolates),Su

illus

gram

ulatus

(3isolates),Su

illus

luteus,

Thelephora

terristris,C

enococcumgeophulum

P.involutus,Hebelom

acrustuliniform

e,Laccaria

laccata

P.involutus,Hebelom

acrustuliniform

e,Laccaria

laccata

Diameter

grow

th

Nutrient

uptake

Nutrient

uptake

Nutrient

uptake

Growth

Growth

Hunt(19

90)

Mitchell,Co

x,Dixon

etal.(19

84)

Mitchell,Co

x,Dixon

etal.(19

84)

Mitchell,Co

x,Dixon

etal.(19

84)

Garbaye

andCh

urin(199

7)

Garbaye

andCh

urin

(199

7)


ReferencesAggangan N C, Lorilla E B, Cruz R E, de La. 1990Screening for effectiveness of four VA mycorrhizal

fungi on Acacia auriculiformis in infertile soilSylvatrop 1987 Publication 12(3&4): 133–140

Ahiabor B D and Hirata H. 1994Characteristic responses of three tropical legumes

to the inoculation of two species of VAM fungi inAndosol soils with different fertilities

Mycorrhiza 5(1): 63–70

Ammani K and Rao A S. 1996Effect of two arbuscular mycorrhizal fungi

Acaulospora spinosa and A. scrobiculata onupland rice varieties

Microbiological Research 151(3): 235–237

Asif M, Khan A G, Khan M A, Kuck C. 1995Growth response of wheat to sheared root vesicular

arbuscular mycorrhizae inoculum under fieldconditions

In Mycorrhizae: biofertilizers for the future, pp. 432–437,edited by A Adholeya and S Singh

New Delhi: Tata Energy Research Institute. 548 pp.[Proceedings of the Third National Conference on

Mycorrhiza, 13–15 March 1995, New Delhi, India]

Boucher A, Dalpe Y, and Charest C. 1999Effect of arbuscular mycorrhizal colonization of

four species of Glomus on physiological re-sponses of maize

Journal of Plant Nutrition 22(4&5): 783–797

Burgess T I, Malajczuk N, and Grove T S. 1993The ability of 16 ectomycorrhizal fungi to increase

growth and phosphorus uptake of Eucalyptusglobulus Labill. and E. diversicolor F. Muell.

Plant and Soil 153(2): 155–164

Diederichs C. 1992Impact of tropical VA mycorrhizae on growth pro-

motion of Cajanus cajan as influence by Psources and P levels

In Proceedings of the International Symposium on Manage-ment of Mycorrhizas in Agriculture, Horticulture andForestry, p. 55, edited by D Jasper

Australia: Australian Institute of Agricultural Sciences.164 pp.

[28 September – 2 October 1992, University of WesternAustralia, Nedlands, Australia]

Douds D D, Galvez L, Becard G, Kapulnik Y. 1998Regulation of arbuscular mycorrhizal development

by plant host and fungus species in alfalfaThe New Phytologist 138(1): 27–35

Ganesan V and Mahadevan A. 1994Effect of mycorrhizal inoculation of cassava, el-

ephant foot yam and taroJournal of Root Crops 20(1): 1–14

Garbaye J and Chorin J L. 1999Growth stimulation of young oak plantations inocu-

lated with ectomycorrhizal fungus Paxillusinvolutus with special reference to summer drought

Forest Ecology and Management 98(3): 221–228

Gaur A, Adholeya A, and Mukerji K G. 1998A comparison of AM fungi inoculants using Capsi-

cum and Polianthes in marginal soil amendedwith organic matter


Gazey C, Abbott L, and Robson A 1992Development of vesicular-arbuscular (VA)

mycorrhizas on clover by three species ofAcaulospora in relation to effects on plant growth

In Proceedings of the International Symposium on Manage-ment of Mycorrhizas in Agriculture, Horticulture andForestry: programme and abstracts, p.32, edited by DJasper

Australia: Australian Institute of Agricultural Sciences.164 pp.

[28 September – 2 October 1992, University of WesternAustralia, Nedlands, Australia]

Ikram A, Mahmud A W, and Othman H. 1993Growth response of Hevea brasiliensis seedling

rootstock to inoculation with vesicular-arbuscular mycorrhizal fungal species in steam-sterilised soil

Journal of Natural Rubber Research 8(3): 231–242

Jaizme-Vega M C and Azcon R. 1995Responses of some tropical and subtropical cul-

tures to endomycorrhizal fungiMycorrhiza 5(3): 213–217

Joner E J and Jakobsen I. 1994Contribution by two arbuscular mycorrhizal fungi

to P uptake by cucumber (Cucumis sativus L.)from (32)P-labelled organic matter during min-eralization in soil

Plant Soil 163(2): 203–209

Lingaraju B S, Sreenivasa M N, and Babalad H B. 1994Response of soybean to VA mycorrhizal fungi at

varied levels of PJournal of Oilseeds Research 11(2): 300–302

Mandal B S and Kaushik J C. 1994Effect of VAM inoculations of growth and develop-

ment of Acacia nilotica (L.)Willd. ex. delAnnals of Biology (Ludhiana) 10(2): 264–267

Mathur N and Vyas A. 1996Relative efficiency of different VAM fungi on growth

and nutrient uptake in Zizyphus mauritianaIndian Journal of Forestry 19(2): 129–131

Mathur N and Vyas A. 1995Production of highly nutritive pods of Cyamopsis

tetragonoloba by VA mycorrhizaeIndian Journal of Experimental Biology 33(6): 464–465

McGraw A C and Schenck N C. 1981The mycorrhizal relationship in tomato as influ-

enced by fungal species, inoculum density, andcultivar

In Proceedings of the Fifth North American Conference onMycorrhizae, p.19, edited by J A Fortin

Canada: University of Laval. 83 pp.[16–21 August 1981, University of Laval, Quebec,

Canada]


Miller D D, Bodmer M, and Schuepp H. 1989Spread of endomycorrhizal colonization and effects

on growth of apple seedlingsThe New Phytologist 111(1): 51–59

Mitchell R J, Cox G S, Dixon R K, Garrett H E, SauderI L. 1984

Inoculation of three Quercus species with elevenisolates of ectomycorrhizal fungi: foliar nutrientcontent and isolate effectiveness

Forest Science 30(3): 563–572

Natarajan K and Reddy S M. 1994Selection of a suitable ectomycorrhizal fungal iso-

late for Pinus patula nurseriesIndian Journal of Microbiology 34(2): 107–112

Nemec S. 1979Growth of Citrus jambhiri as affected by single and

combined incocula of three Glomus speciesIn Proceedings of the Fourth North American Conference on

MycorrhizaColorado: Colorado State University[24–28 June 1979, Colorado State University, Fort

Collins, Colorado, USA]

Obura R K, Skipper H D, and Wagner S C. 1987Effect of VAM fungi on cowpea under two levels of

soil phosphorusIn Proceedings of the Seventh North American Conference

on Mycorrhiza, p. 54, edited by D M Sylvia, L LHung, and J H Graham

Florida: University of Florida. 364 pp.[3–8 May 1987, Gainesville, Florida]

Parlade J, Alvarez I F, and Pera J. 1996Ability of native ectomycorrhizal fungi from north-

ern Spain to colonize Douglas-fir and other in-troduced conifers


Ponton F, Piche Y, Parent S, Caron M. 1990Use of vesicular-arbuscular mycorrhizae in Boston

fern production. II. evaluation of four inoculaHorticultural Science 25(4): 416–419

Rajan S K, Reddy B J D, and Bhagyaraj D J. 2000Screening of arbuscular mycorrhizal fungi for their

symbiotic efficiency with Tectona grandisForest Ecology and Management 126(2): 91–95

Ramana B V and Babu R S H. 1999Response of onion (Allium cepa) to inoculation of

VAM fungi at different levels of phosphorusIn Proceedings of the National Conference on Mycorrhiza.

Section 2 (Poster): Plant growth responses andmycorrhizal dependency, edited by S Singh

India: Bhopal[5–7 March 1999, Bhopal, India]

Rao V U and Rao A S. 1996Response of blackgram (Phaseolus mungo) and

greengram (P. radiatus) to inoculation with ve-sicular-arbuscular mycorrhizal fungus underdifferent sources of phosphorus fertilizer

Indian Journal of Agricultural Sciences 66(1): 613–616

Rathore V P and Singh H P. 1995Influence of vesicular-arbuscular mycorrhizal

fungi and phosphate on maizeJournal of the Indian Society of Soil Science 43(2): 207–210

Reid R K, Parker W C, Mitchell R J, Garrett L C,Hundersen G S, Cox G S. 1988

Vesicular arbuscular mycorrhizal inferences on growthand aluminium tolerance of sugar maple seedlings

University of British Colombia Publication, pp. 306–315

Rodriguez G A M, Hurtado M, and Sanchez de PragerM. 1995

Inoculation of granadilla Passiflora ligularis L.with VAM

Acta Agronomica, Universidad Nacional de Colombia45(1): 89–98

Secilia J and Bagyaraj D J. 1994Evaluation and first-year field testing of efficient

vesicular arbuscular mycorrhizal fungi for in-oculation of wetland rice seedlings

World Journal of Microbiology and Biotechnology 10(4):381–384

Shaw C G III and Molina R. 1979Ectomycorrhizal inoculation of containerized sitka

spruce seedlingsIn Proceedings of the Fourth North American Conference on



Sivaprasad P, Sulochana K K, and Nair S K. 1990Comparative efficiency of different VA mycorrhizal

fungi on cassava (Manihot esculenta Crantz)Journal of Root Crops 16(1): 39–40

Struble J E, Skipper H D, and Smith G W. 1979Effects of soil pH on the growth response of corn to

VA mycorrhizaeIn Proceedings of the Fourth North American Conference on



Sumana D A and Bagyaraj D J. 1999Selection of efficient VA mycorrhizal fungi for in-

oculating neemIn Proceedings of the National Conference on Mycorrhiza.

Section 2: Association and distribution of mycorrhizalfungi under varying edapho-climatic conditions, ed-ited by S Singh

[5–7 March 1999, Bhopal, India]

Taber R A, Smith O D, Smith D H, Pettit R E, Neck JS, Rajapakse S, Harman W F. 1987

Growth response of Arachis hypogaea to field in-oculation with vesicular-arbuscularendomycorrhizal fungi

In Proceedings of the Seventh North American Conferenceon Mycorrhiza, p. 38, edited by D M Sylvia, L LHung, and J H Graham



Taube-Baab H and Baltruschat H. 1996Productivity of strawberry (fragaria × ananassa

Duch.) plants following inoculation with VAmycorrhizal fungi

Erwerbsobstbau 38(5): 144–147

Tewari L, Johri B N, and Tandon S M.1993Host genotype dependency and growth enhancing

ability of VA-mycorrhizal fungi for Eleusinecoracana (finger millet)

World Journal of Microbiology Biotechnology 9(2): 191–195

Thomson J, Matthes-Sears U, and Peterson R L. 1990Effects of seed provenance and mycorrhizal fungi

on early seedling growth in Picea marianaCanadian Journal of Forest Research 20(11): 1739–1745

Torres P and Honrubia M. 1994Inoculation of containerized Pinus halepensis

(Miller) seedlings with basidiospores ofPisolithus arhizus (Pers.) Rauschert,Rhizopogon roseolus (Corda)Th. M. Fr. andSuillus collinitus (Fr.) O.Kuntze

Annales des Sciences Forestieres 51(5): 521–528

Tyminska A, Tacon F Le, and Chadoeuf J. 1986Effect of three ectomycorrhizal fungi on growth

and phosphorus uptake of Pinus silvestris seed-lings at increasing phosphorus levels

Canadian Journal of Botany 64(11): 2753–2757

Vasuvat Y, Wadisirisak P, Toomsen B, Nopamornbodi O,Thamsurakul S, Thananusont V, Boonkerd N. 1987

Interaction between mycorrhizal fungi and cowpearhizobia on peanut cultivar Tainan-9 in Thailand

In Proceedings of the Seventh North American Conferenceon Mycorrhiza, p. 61, edited by D M Sylvia, L LHung, and J H Graham


Vijaykumar B S and Bhiravamurthy P V. 1999Selection of efficient indigenous local VAM fungi

for improving the groundnut crop in semi-aridtropical soils of Anantapur district, AndhraPradesh

In Proceedings of the National Conference on Mycorrhiza.Section 2 (Poster): Plant growth responses andmycorrhizal dependency

[5–7 March 1999, Bhopal, India]

Zhu H and Navratil S.1987Effect of seed source on growth and ectomycorrhizal

formation of tamarack seedlingsIn Proceedings of the Seventh North American Conference

on Mycorrhiza, p.77, edited by D M Sylvia, L LHung, and J H Graham

Florida: University of Florida[3–8 May 1987, Gainesville, Florida]

Research findings

Morphological and cytochemical study of the pelotonic and non-pelotonichyphae of the orchid Spathoglottis plicata

S Senthilkumar* and S John BritooDepartment of Botany, St. Joseph’s College, Tiruchchirappalli – 620 002, India

IntroductionOrchids have a special type of endomycorrhizae inwhich the fungal hyphae produce very characteristicpelotons (coiled balls) in the host cortical cells(Senthilkumar and Krishnamurthy 1998). Themycorrhizal association with orchids has been thesubject of intensive study, yet the nature of the rela-tionship remains obscure. Peterson, Mueller, andEnglander (1980) described a relationship involving atleast two different fungi (a basidiomycete and an asco-mycete). Later, Couture, Fortin, and Dalpe (1983)obtained isolates of a sterile fungus and Oidiodendrongriseum from the roots of Vaccinium. Resolution ofthese differences is hampered by the inability to isolateand identity some of the fungi (Muller, Tessier, andEnglader1985) and by the inability to differentiate thefungi by cytochemical procedures. To overcome thelatter problem, we have used a variety of cytochemicaland cytoenzymological procedures for direct differen-tiation of the fungi in roots. We describe here theprocedure for pelotonic and non-pelotonic hyphae inthe roots of Spathoglottis plicata.

Materials and methodsSpathoglottis plicata, an ornamental ground orchidcommonly available in several parts of Sri Lankacontaining the natural inoculum of Rhizoctonia wascollected from the botanical garden of PeradaniyaUniversity, Sri Lanka. The mycorrhizal roots werecut into small pieces, washed thoroughly in water,and fixed in FAA (formalin – acetic acid – alcohol).Free-hand sections were made for several cyto-chemical procedures to understand the changesnoticed in hyphae and in the host cells(Krishnamurthy 1998) (Table 1).

Results and discussionInfection of the root is always by hyphae throughroot hairs, which are commonly at root tips. Usu-ally, only a single hypha enters into each root hair,but occasionally two or rarely more than two findentry into a single hair. Fungal entry is always con-fined to the part of the root that has root hairs, andno other region of the root is infected (Burgeff1959). Entry through root hairs as well as

*E-mail: [email protected]


rhizodermis has also been reported by a few investi-gators (Peterson and Farquhar 1994; Senthilkumarand Krishnamurthy 1998). At the point of entry,no special features can be seen in the hypha. Theenzymatic dissolution of the root hair wall by thefungus probably facilities its entry.

Entry into the cortex is always through the pas-sage cells of the exodermis, and no instance of fun-gal mycelium in the thick-walled cells of theexodermis has been noticed. From this location,the hyphae again intracellularly enter into the cor-tex proper and some of the branches get into thecortical parenchyma cells for forming pelotons.

However, subsequent hyphae settle down gradu-ally in the middle regions without forming pelotons.These cortical cells often show only hyphal filamentsand no hyphal clumps or pelotons; in other words,the capacity to form pelotons is restricted to onlythe orchid mycorrhizae. As in many previouslystudied orchids, colonization was restricted to thecortex only. Penetration of hyphae into endodermisand central cylinder, as observed by Ruinen (1953)in a species of Dendrobium, was not observed in thepresent orchid.

However, as infection is a continuous processand all cells of the cortex are not colonized at thesame time, there is often a mix-up of pelotons andpelotonic hyphal colonization across the entire cor-tex (Figure 1). One significant event in the orchid–mycorrhizal association is the lysis of the pelotons(Peterson and Currah 1990; Senthilkumar andKrishnamurthy 1998). Digestion was also observedrandomly in the cortex and not in a defined zone of

cells, designated as the ‘digestion layers’, whichhave been recognized in the inner cortex of someorchid species (Hadley 1982; Hadley andWilliamson 1971). But interestingly, the digestionof non-pelotonic mycorrhizal elements was notnoticed in the cortical cells. Fusconi and Bonfante-Fasolo (1984) described the mycorrhizae formedbetween Arbutus unedo and an unknown ascomyc-ete symbiont and found a typical hartig net, intrac-ellular colonization, and a thin fungal mantle.

A number of investigators have carried outdetailed structural and cytochemical studies on the

Table 1 Cytochemical procedures

Procedures employed and Substances to be localized,treatment time colour to be obtained Rationale

Acid fuchsin method Protein: Reddish pink Aldehydes of proteins react with acid fuchsin(1% in distilled water) (Robinow and Marak 1966) and get stained

Alcian blue (3%) solution in 0.1N Carboxylated polysaccharides: Negatively charged carboxyl group of uronic acidssodium acetate buffer, pH 5.6 blue (Gahan 1984) of pectins bind to the cationic alcian blue by salt

linkages at the isoindole conferred cationic sites

Chlorazol black E Cellulose: black to bluish black Relation not clear(in 1% methyl cellosolve 5 min) (Krishnamurthy 1998)

Commassie Brilliant Blue Total proteins: blue Yields a stereo-chromatic reaction with(0.02% in Clarke�s solution 15 min) (Krishnamurthy 1998) all proteins

Fast green FCF (fast green FCF in 50 ml of Basic protein and nuclei: green Deamination/acetylation/0.1 M phosphate buffer pH 8.0) 30 min (Prento and Lyon 1973; Gahan 1984) trypsin extraction

Sudan black E Lipids: black (Krishnamurthy 1998) This dye gets selectively soluble in tissue lipids,thereby staining the complex black to blue black

Toluidine blue O (TBO) Carboxylated polysaccharides: TBO is a meta-chromatic dye. Highly(0.5% in acetate buffer, pH 4.4) pink to reddish pink (Mc Cully 1966) polymerized lignin shows orthochromatic blue

colour and less polymerized lignin bluish colour

Figure 1 Cross section of rootstained with Alcain blue to showthe pelotons (single arrow) andnon-pelotonic hyphae (doublearrow) (× 150).


pelotons (Peterson and Currah 1990; Senthilkumarand Krishnamurthy, 1998). A number of theseobservations are confined to orchid mycorrhizasand non-pelotonic hyphae, as is also the case in thepresent study. The orchid mycorrhizae are fullyfilled with cytoplasm and get stained prominentlywith fast green, indicating their high histone con-tent. The cytoplasm is rich in basic proteins andinsoluble polyaccharides (Figure 2), and has fairlygood amounts of lipids and cellulose. InPlatanthera, Richardson, Petewrso, and Currah(1992) observed lipids polyphosphates in the hy-phae in the initial stage of lysis. Non-pelotonichyphae remained intact whereas pelotonic hyphaeunderwent digestion (Figure 3). Intense activity ofcarboxylated polysaccharides was found in the cellwall (Figure 2), although activity could also bedetected to some extent in other regions. The

present study indicated that there are goodamounts of basic proteins, lipids, and cellulose,especially basic protein in the cell wall and cyto-plasm of fresh non-pelotonic filaments. Barroso,Chaves, and Pais (1986) reported that there are anumber of steroidal substances in the infected cellsof Ophrys lutea. They may also perhaps be responsi-ble for the positivity that was observed for lipidstains used in the present study. There is also amoderate activity of histones and poor activity ofpolysaccharides in the non-pelotonic hyphae. Non-pelotonic hyphae are separated from the lysingpeloton by the presence of phenols in their walls.This is one of the reasons for the high activity ofphenols in their walls.

Among these associations, one of the hyphae(orchid mycorrhizae) enters into root hairs andstarts producing shorter hyphal cells. It may remainunbranched or often undergo branching; the twobranched hyphae are seen parallel to each other inhost root hair. One or both of these hyphalbranches soon produce the so-calledchlamydospores (Saksena and Vaartaja 1961).These chlamydospores are either terminally pro-duced in these hyphae or are intercalary and maybe produced singly or more commonly in chains,resulting in beaded or moniliform appearance (Fig-ure 4). Chlamydospores are pyriform, oval, spheri-cal, or club-shaped, the terminal ones oftenshowing a beak. The hyphae producing thechlamydospores are rich in total proteins, carboxy-lated polysaccharides, and phenols. Sneh,Burpee, and Ogoshi (1991) and Senthilkumar,Krishnamurthy, and Vengadeshwari (1998) re-ported that these types of chlamydospores are

Figure 2 Cross sections of root indicating the presenceof polysaccarides in the pelotons and colouration indi-cating the presence of phenols with TBO (× 650).

Figure 3 The cortical cellshowing lysing pelotons(arrow) with non-pelotonichyphae (double arrow).

Figure 4 Chains of chlamydospores in root hair about tobe released (arrow) (× 650).


occasionally produced in epidermal cells and roothairs of the host. To the best of our knowledge,there is no report of these two types of associationin the orchid Spathoglottis plicata, and there is nosatisfactory explanation of the role of non-pelotonicmycorrhizae.

AcknowledgementsThe authors express gratitude to Prof. S A Kulasoorya,Head, Department of Botany, Peradaniya University,Sri Lanka, for granting permission to perform theexperiments and Director, Peradaniya BotanicalGarden, for orchid root samples.

ReferencesBarroso J, Chaves H, and Pais M S. 1986Ultrasturctural, cytochemical and biochemical

aspects related to the formation of Ophrys luteaendomycorrhizae

In Physiological and Genetical Aspects of Mycorrhizae, pp.265–269, edited by V Gianinazzi-Pearson and SGianinazzi

[Proceedings of the First European Symposium onMycorrhizae, Dijon]

Burgeff H. 1959Mycorrhiza of orchidsIn The Orchids: scientific survey, pp. 361–395, edited by

C L WithnerNew York: Ronald Press

Couture M, Fortin J A, and Dalpe Y. 1983Oidiodendron griseum Robak: an endophyte of

ericoid mycorrhiza in Vaccinium sp.The New Phytologist 95: 375–380

Fusconi A and Bonfante-Fasolo P. 1984Ultrastructural aspects of host-endophyte relation-

ships in Arbutus unedo L. mycorrhizasThe New Phytologist 96: 379–410

Gahan P B. 1984Plant Histochemistry and Cytochemistry: introductionFlorida: Academic Press

Haldley G. 1982Orchid mycorrhizaIn Orchid Biology: reviews and perspectives II, pp. 83–118,

edited by J ArdittiIthaca, New York: Cornell University Press

Hadley G and Williamson B. 1971Analysis of the post infection growth stimulus in

orchid mycorrhizaThe New Phytologist 70: 445–455

Krishnamurthy K V. 1998Cell Wall CytochemistryUSA: CRC Press 325 pp.

Mc Cully M E. 1966Histological studies on the genus Fucus. I. Light

microscopy of the mature vegetative plantProtoplasma 62: 287–305

Muller W C, Tessier B J, and Englander L. 1985Immunocytochemical detection of fungi in the

roots of RhododendronCanadian Journal of Botany 64: 718–723

Peterson T A, Mueller W C, and Englander L. 1980Anatomy and ultrastructure of a Rhododendron

root–fungus associationCanadian Journal of Botany 58: 2421–2433

Peterson R L and Currah R S. 1990Synthesis of mycorrhizae between protocorms of

Goodyera repens (Orchidaceae) andCeratobasidium cereale

Canadian Journal of Botany 68: 1117–1125

Peterson R L and Farquhar M L. 1994Mycorrhizas – integrated development between

roots and fungiMycologia 86: 311–326

Prento P and Lyon H. 1973Nucleoprotein staining: an analysis of a standard

trichloroacetic acid – fast green FCF procedureHistochemical Journal 5: 493–501

Richardson K A, Petewrso R L, and Currah R S. 1992Seed reserves and early symbiotic protocorm develop-

ment of Platanthera hyperborea (Oprchidaceae)Canadian Journal of Botany 70: 291–300

Robinow C F and Marak J. 1966A fibre apparatus in the nucleus of the yeast cellJournal of Cell Biology 29: 129–151

Ruinen J. 1953Epiphytosis: a second view on epiphytismAnn. Bogor. 1: 101–157

Saksena H K and Vaartaja O. 1961Taxonomy, morphology and pathogenicity of

Rhizoctonia species from forest nurseriesCanadian Journal of Botany 39: 627–646

Senthilkumar S and Krishnamurthy K V. 1998A cytochemical study on the mycorrhizae of

Spathoglottis plicataBiologia Plantarum 41: 111–119

Senthilkumar S, Krishnamurthy K V, and VengadeshwariG N. 1998

Studies on the mycorrhizal association of the orna-mental orchid, Papilonanthe subulata

Philippines Journal of Science 127: 189–199

Sneh B, Burpee L, and Ogoshi A. 1991Identification of Rhizotonia Species, pp. 1–4USA: Academic Press. 133 pp.


IntroductionVesicular–arbuscular mycorrhizal (VAM) fungiform obligate symbiotic associations with the rootsand other underground parts of most plants. Theirassociation with plant roots indirectly resists pen-etration by nematodes, thus helping the plant in itsgrowth and production. It is well known that VAmycorrhizal fungi improve the growth of plants byproviding higher absorptive surface compared withroot hairs, and thus help in the absorption of rela-tively immobile ions in soil, such as phosphorus,copper, and zinc (Bagyaraj, 1992). Continued ef-forts are on to develop mass production technolo-gies for VAM fungi. Hence, using various hosts, astudy was undertaken to compare the mass multi-plication of VAM.

Materials and methodsDifferent host crops (11) were grown in pots con-taining sterilized soil, and a culture of VAM

(Glomus fasciculatum) was added at 10 g/kg soil formultiplication. After one and two months, spores insoil and root colonization were recorded to find thesuitable preferred host for VAM multiplication.

Results and discussionPearl millet (Cumbu) (Pennisetum typhoides) wasfound to be the best host—the spore populationwas 460 per 200 ml soil after one month (Table 1).This was followed by bhendi with a spore popula-tion of 268.8 per 200 ml soil after one month. Incotton, spore count reached 312.9 per 200 ml afterthe second month. The spore population was morethan 100 in 200 ml soil of cowpea, maize, and redgram whereas in all other hosts, the count was lessthan 100. Sieverding and Leihner (1984) have re-ported that graminaceous and leguminous cropsgenerally tend to increase VAM population whereasmycotrophic plants decrease the population ofVAM fungi.

Mass production of vesicular�arbuscular mycorrhizae using different hosts

R Sundara Babu, K Poornima, and N SugunaDepartment of Plant Nematology, Tamil Nadu Agricultural University, Coimbatore – 641 003, India

Table 1 Mass multiplication of VAM: host preference

Host Shoot length Shoot weight Root length Root weight Spore count/ VAM colonization(m) (g) (cm) (g) 200 ml soil (%)

TomatoMonth I 61.8 19.9 17.8 6.6 25.9 20.0Month II 79.3 43.2 14.2 15.3 87.1 43.9

CumbuMonth I 56.9 1.7 14.2 0.2 460.0 71.2Month II 73.8 9.5 18.8 1.1 86.7 80.3

CottonMonth I 37.2 0.6 11.0 0.4 60.1 71.0Month II 38.8 4.2 12.1 1.1 312.9 60.5

BhendiMonth I 27.6 6.0 13.3 1.3 268.8 74.5Month II 44.6 12.0 13.3 2.4 222.9 70.0

BrinjalMonth I 20.3 3.8 13.7 2.1 85.3 31.0Month II 27.0 8.3 17.4 2.4 41.4 36.0

CowpeaMonth I 39.1 6.7 14.7 0.7 167.0 67.0Month II 86.5 9.8 11.8 1.5 199.0 70.0

Green gramMonth I 37.7 2.4 10.1 0.6 113.0 68.0Month II 34.9 3.1 27.7 4.3 73.0 49.0

MaizeMonth I 65.0 8.6 74.0 18.6 170.0 68.0Month II 92.1 17.7 44.2 18.9 143.0 60.8

Red gramMonth I 37.7 1.5 12.5 0.6 152.4 56.2Month II 42.1 3.2 3.4 1.1 130.6 60.8

Black gramMonth I 30.0 3.1 12.5 0.4 79.4 103.3Month II 11.0 6.4 15.9 5.1 88.9 46.8


AcknowledgementsThe authors are grateful to ICAR for financial as-sistance of the ad-hoc scheme ‘Mass production ofVAM using differential hosts’.

ReferencesBagyaraj D J. 1992Vesicular–arbuscular mycorrhiza: application in

agricultureMethods in Microbiology 24: 359–374

Severding E and Leiher D E. 1984Influence of crop rotation and intercropping of

cassava with legumes on VA mycorrhizal sym-biosis of cassava

Plant Soil 80: 143–146

Root trainers are being effectively used to raiseseedlings in forest nurseries. Different grades ofpotting mixture are used as potting media in roottrainers (Mishra and Emmanuel 1997). In mostcases, compost is a suitable substrate for raisingseedlings in root trainers. It has been observed thatcompost does not contain an effective VAM popu-lation. However, a very sparse population of sporeshas been recorded in compost, which mostly occuras contaminants through soil, root bits, etc. More-over, pure compost has an acidic pH, which can beamended by using sand. The role of VAM (vesicu-lar–arbuscular mycorrhiza) in improving the qualityand survival of forest tree seedlings is well docu-mented. There is considerable variability in VAMdevelopment in different grades of potting mixture.An experiment was conducted to study VAM in theroot trainer by using Albizia procera as the test spe-cies. The combination of treatments is as follows.1 Pure compost, pH 5.82 Compost + sand (80:20), pH 6.43 Compost + sand (50:50), pH 6.44 Sand, pH 6.8

The capacity of root trainers was 300 cc. Thecompost contained Glomus, Scutellospora, andGigaspora in very low quantity. When the compostis diluted by using sand, the population ofpropagules of VAM is reduced considerably. Seedswere sown in the root trainer after proper treat-ments (Rahangdale and Gupta 1998). The VAMculture containing root bits and soil was appliedafter initiation of seed germination (after 10 days).The inoculation was made by using the mix VAMculture prepared from A. procera by using Panicummaximum as the trap plant. The inoculum was ap-plied at the rate of 25 g/root trainer with a capacityof 300 cc having 250 infective propagules. Infectivepropagules included Gigaspora spp., Scutellosporaspp., Glomus mosseae, G. intraradices, andAcaulospora scrobiculata. VAM colonization in roots

A note on vesicular�arbuscular mycorrhiza development in Albiziaprocera in root trainers

M G Goswani and JamaluddinTropical Forest Research Institute, P.O. - R.F.R.C., Jabalpur – 482 021

of A. procera was obtained by using the techniquesof Phillips and Hayman (1970). The experimentwas conducted for three months.

The compost contained a sparse population ofVAM, particularly Gigaspora, Scutellospora, andGlomus mosseae. After dilution with sand, the popula-tion is reduced considerably. The pH of the pottingmedia was also measured. After treatment with VAM,the spore number of VAM fungi increased in thepotting mixture containing compost + sand (80:20,50:50) and pure compost. However, spore develop-ment is low in pure sand. The per cent colonizationof root by VAM was higher in plants grown in pottingmedia having compost and sand (80:20) and purecompost. Spore population was lower in pure com-post with pH 5.8. A similar trend was noticed byRaghvendra and Lakshman (1995). The height ofplants varied from 31 cm to 15 cm in different treat-ments (Table 1). The height growth of A. procera was31 cm, 27 cm, and 21 cm in pure compost, compost

Table 1 VAM status and seedlings growth of A. procera in roottrainers in different combinations of compost after 4 months

Shoot No. No. of

Infection height of spores/

Treatment (%) (cm) nodules 100 g pH

Pure compost 45 31.00 20.00 180.00 5.8

Compost + 50 27.00 34.00 200.00 6.4

sand (80:20)

Compost + 40 21.5 33.00 260.00 6.4

sand (50:50)

Sand 40 20.00 18.00 189.00 6.8

SEM +2.39 +2.54 +4.21 +18.05

CD at 5% 9.39 9.95 16.53 70.00


+ sand (80:20), and compost + sand (50:50) respec-tively. Verma and Jamaluddin (1994) reported that amixture of Glomus mosseae, G. fasciculatum, G.etunicatum, and G. aggregatum enhanced growth ofAcacia nilotica seedlings under different moistureregimes. In the present study, VAM inocula having aconsortium of VAM fungi, namely Glomus mosseae, G.aggregatum, G. intraradices, Scutellospora, Gigaspora,and Acaulospora scrobiculata, colonized successfully indifferent potting media. However, mycorrhizal infec-tion and spore number were low in pure sand.Rahangdale and Gupta (1998) noticed only a 26.6%infection of VAM in A. procera. However, in thepresent study, VAM colonization reached a level of50%. Root nodule development was more whenplants were grown in compost + sand (80:20 and50:50). It is concluded that the potting mixture withpure compost or its amendment with sand requireinoculation with VAM consortium to enhance thegrowth of A. procera in root trainers. The populationof the VAM also multiplies significantly in such pot-ting media.

ReferencesMishra D K and Emmanuel C J S K. 1997Use of root trainers in forestry: a cautious approachMy Forest 33(4): 583–588

Phillips J M and Hayman D S. 1970Improved procedures for cleaning roots and stain-

ing vesicular–arbuscular mycorrhizal fungi forrapid assessment of infection

Transactions of the British Mycological Society 55: 158–161

Raghvendra S and Lakshman H G. 1995Effect of different soil pH values on vesicular–

arbuscular mycorrhizal associated tree speciesIn Mycorrhizae: biofertilizers for the future, pp. 510–515,

edited by A Adholeya and Sujan Singh[Proceedings of the Third National Conference on

Mycorrhiza, 13–15 March 1995, New Delhi, India]

Rahangdale R and Gupta N. 1998Selection of VAM inoculates for some forest tree

speciesIndian Forester 124(5): 331–341

Verma R K and Jamaluddin. 1994Effect of VAM fungi on growth and survival of Aca-

cia nilotica seedlings under different moistureregimes

Proceedings of National Academy of Science India 64(B)II:205–210

Comparison of various techniques forrecording AM colonization

The extent of AM colonization was assessed byGange A C, Bower E, Stagg P G, Aplin D M,Gillam A E, Bracken M (1999) in 10 field-collectedplant species—three annual forbs, three perennialforbs, three perennial grasses, and one annual grass(The New Phytologist 142(1): 123–132). The rootsystem of each plant was split into four portions,and the mycorrhizal structures were revealed usingfour methods, i.e. epifluorescence microscopy (un-der which only arbuscules are generally visible) andthree stains (chlorazol black E, acid fuchsin, andtrypan blue) to evaluate the efficiency of eachmethod and compare results by each method understandard laboratory conditions. The recorded colo-nization levels of arbuscules, total arbuscularmycorrhizal fungal material, and total fungal(arbuscular mycorrhizal + non-arbuscularmycorrhizal) material differed significantly withdifferent visualization methods in a number of spe-cies. However, there were also interactions betweena stain and plant species, indicating that the per-formance of a stain is dependent on the plant spe-cies being examined. In some cases (e.g. Plantagolanceolata), each visualization method produced thesame level of colonization, whereas in others (e.g.Dactylis glomerata), each method gave a differentresult. The findings, therefore, suggest that the

level of mycorrhizal colonization recorded in anyparticular plant species at a particular time is de-pendent on the technique employed.

Molecular methods for determiningintra- and interspecific variations inectomycorrhizal fungi

Molecular techniques were employed by BonfanteP, Lanfranco L, Cometti V, Genre A (1997) ongenomes of 13 Suillus isolates from Mediterraneanand Alpine regions by using a range of PCR-basedtechniques (Microbiological Research 152(3): 287–292). Amplification of the ITS (inter-transcribedspacers) region gave fragments of the same lengthirrespective of the species, whereas RFLP (restric-tion fragment length polymorphism) revealedinterspecific variability. Random oligonucleotidesfor RAPD (random amplified polymorphic DNA)amplification and primers for microsatellites led topolymorphic fingerprints, revealing intraspecificdifferences. In addition, microsatellite finger-printing allowed separation of strains with differentsymbiotic capabilities (e.g. Suillus collinitus strains24 and 31) as well as strains originating from thesame zone. The studies thus suggest that a range ofmolecular probes used on the same isolates canprovide both specific fingerprints for selected fun-gal strains and tools to investigate the physiologicalcharacteristics of ectomycorrhizal fungi.

New approaches


Mycorrhiza in sustainable agriculture

Reena Singh and Alok AdholeyaCentre for Mycorrhizal Research, Tata Energy Research Institute, Darbari Seth Block, Habitat Centre,Lodhi Road, New Delhi – 110 003, India

attributed to an intensification-induced degradationprocesses (Pingali, Hossain, and Gerpacho, 1997).

Such degradation of arable land is a majorthreat worldwide to the sustainability of agricul-tural production. During the last 40 years, nearlyone-third of the world’s arable land has been lostby erosion and continues to be lost at a rate ofmore than 10 million ha per year (Pimental,Harvey, Resosudarmo, et al. 1995). For example,in India, 2400 million tonnes of silt is estimated tobe carried away through the rivers every year; fur-thermore, 4.5 million ha is affected by salinizationand 6 million ha by waterlogging due to agricul-tural (mis-)management (Pingali, Hossain, andGerpacio 1997). A more holistic, long-term ap-proach to food production in sustainableagroecosystems is therefore urgently needed.The main goal of such an approach is to maintaineconomic crop production levels in a sustainable,ecologically sound way. To attain this goal, farmingstrategies have to be adopted that exploit and en-hance inherent resources of the natural environment,such as soil fertility, biological nitrogen fixation, bio-logical control of diseases, etc., rather than relying onhigh input of costly external resources.

The symbiosis of plants with AMF (arbuscularmycorrhizal fungi) is the most widespreadmutualistic symbiosis in natural ecosystems, whichhas been estimated to occur on land growing wheatand pulses and almost all other important agricul-tural crops. It is well documented that mycorrhizalplants show improved growth, health, and resist-ance to abiotic and biotic stress as compared withnon-mycorrhizal controls. These beneficial effectsof AMF for the plants are expressed most understresses such as nutrient limitation, drought, orheavy metal pollution. The AM symbiosis has alsobeen shown to contribute substantially to soil con-servation via its role in the formation of water-sta-ble soil aggregates by extraradical mycelia and thebacteria associated with it. These aggregates arecrucial for creating and maintaining amacroporous, water-permeable soil structure,which is a prerequisite for erosion resistance andaerobic microbial activity in the rhizosphere, neces-sary for efficient nutrient cycling. Due to thesefunctional attributes, AM symbiosis is of particularrelevance for low-input agriculture in the third

Centre for Mycorrhizal Culture Collection

The word ‘sustain’, from the Latin sustinere (sub:‘from below’ and tenere: ‘to hold’), means ‘to keep inexistence or maintain’, and implies long-term supportor permanence. As regards agriculture, ‘sustainable’describes farming systems that are ‘capable of main-taining their productivity and usefulness to societyindefinitely. Such systems must be resource-conserving, socially supportive, commercially com-petitive, and environmentally sound.’ (Duesterhaus,1990). Sustainable agriculture was addressed by theCongress in the 1990 Farm Bill (FACTA [Food,Agriculture, Conservation and Trade Act] of 1990)under which the term ‘sustainable agriculture’ means‘an integrated system of plant and animal productionpractices having a site-specific application that will,over the long termn satisfy human food and fiber needsn enhance environmental quality and the natural

resource base upon which the agriculturaleconomy depends

n make the most efficient use of nonrenewableresources and on-farm resources and integrate,where appropriate, natural

n biological cycles and controlsn sustain the economic viability of farm operationsn enhance the quality of life for farmers and soci-

ety as a whole.’

The strategy being adopted in India as well asin other Asian countries essentially relies on twointerrelated actions: first, a dramatic intensificationof input of external resources such as water (irriga-tion systems), fertilizer, and a plethora of pesticidesand other agrochemicals, and second, the introduc-tion of modern, high-yielding, but generally alsohighly demanding crop varieties which are oftenplanted in continuous, uniform monocropping sys-tems. It is now clear that agricultural ecosystemssubjected to continuous high intensification invari-ably exhibit degradation trends over the long term.This is mainly due to adverse effects on the soilstructure, health, and fertility, along with an in-crease in number and intensity of pests. A recentbroad survey of the IRRI (International Rice Re-search Institute) covering several intensively man-aged agricultural areas in India and the Philippinesrevealed that crop yields remained stagnant oreven declined during the last decade and this was


world as well as for the recultivation and manage-ment of degraded soils and wasteland. It is also ofgeneral relevance to the global endeavour to de-velop more sustainable forms of agriculture (forreviews, see Bethlenfalvay and Linderman 1992;Pfleger and Linderman 1994; Sieverding 1991;Smith and Read 1997).

The CMR (Centre for Mycorrhizal Research)group at TERI is actively involved in the activitiesrelated to sustainable agriculture. An integrated useof organic manure and mycorrhizal biofertilizerworks wonders in the agricultural fields. The CMRhas specific mycorrhizal fungi for many kinds ofplants: vegetables (onions, potatoes, etc.), foddercrops (barseem, lucerne, sorghum, etc.), flowers(marigold, gladiolus, aster, etc.), trees (eucalyptus,poplar, etc.). Use of mycorrhizal fungi has beenfound to increase yields by 30%–50%.

Without doubt, an improved understandingand management of the symbiosis of plants withAMF in agroecosystems ultimately has a large so-cial and environmental impact, particularly in low-input sustainable agriculture and in tropicalagrosystems. It is certainly of great importance forIndia, where nearly one half of the districts havebeen classified as being highly deficient in plant-available phosphorus (Pingali, Hossain, andGerpacio 1997).

ReferencesBethlenfalvay G J and Linderman R G. (Eds.) 1992Mycorrhizae in sustainable agricultureMadison, WI: American Society of Agronomy, Inc. 124 pp.[ASA Special Publication No. 54]

Duesterhaus R. 1990Sustainability’s promiseJournal of Soil and Water Conservation 45: 4

Food, Agriculture, Conservation, and Trade Act(FACTA). 1990

Public Law 101-624, Title XVI, Subtitle A, Section1603

Washington, DC: Government Printing Office[NAL Call = KF1692.A31 1990]

Pfleger F L and Linderman R G. 1994Mycorrhiza and Plant HealthSt. Paul, Minnesota: The American Phytopathological

Society Press

Pimentel D, Harvey C, Resosudarmo P, Sinclair K,Kurz D, McNair M, Crist S, Shpritz L, Fitton L,Saffouri R, Blair R. 1995

Environmental and economic costs of soil erosionand conservation benefits

Science 267: 1117–1123

Pingali P L, Hossain M, and Gerpacio R V. 1997Asian rice bowls: the returning crisis?CAB International. 341 pp.

Sieverding E. 1991Vesicular-arbuscular Mycorrhiza Management in

Tropical AgrosystemsGermany, Eschborn: Technical Corporation. 371 pp.

Smith S E and Read D J. 1997Mycorrhizal SymbiosisLondon: Academic Press. 605 pp.

FORM IV(as per Rule 8)

Statements regarding ownership and other particulars about Newsletter: Mycorrhiza News

1 Place of publication New Delhi2 Periodicity of publication Quarterly3 Printer’s name Dr R K Pachauri

Whether citizen of India YesAddress Tata Energy Research Institute,

Darbari Seth Block, Habitat Place,Lodhi Road, New Delhi- 110 003

4 Publisher’s name Dr R K PachauriWhether citizen of India YesAddress Tata Energy Research Institute,


5 Editor’s name Dr Alok AdholeyaWhether citizen of India YesAddress Tata Energy Research Institute,


I, R K Pachauri, hereby declare that the particulars given above are true to the best of my knowledge and belief.

Signature of Publisher(Sd/-)

Date: April 2001 Dr R K Pachauri


Recent references

Latest additions to the Network’s database on mycorrhiza are published here for information of the mem-bers. The Mycorrhiza Network will be pleased to supply any of the available documents to bonafide re-searchers at a nominal charge.

This list consists of papers from the following journals, which are arranged in alphabetical order.

n Annals of Forest Sciencen Biology and Fertility of Soilsn Biotechnology Lettersn Ecology Lettersn FEMS Microbiology Ecologyn Fungal Associationsn Greece Agrochimican Journal of Experimental Botanyn Journal of Plant Growth Regulation

n Naturwissenschaftenn Nova Hedwigian Mycological Researchn Physiological and Molecular Plant Pathologyn Phytochemistryn Plant Biologyn Potato Researchn The New Phytologist

Copies of papers published by mycorrhizologists during this quarter may please be sent to the Network forinclusion in the next issue.

Name of the author(s) and yearof publication

Title of the article, name of the journal, volume no., issue no., page nos[*address of the corresponding author]

Effects of endomycorrhizal development and light regimes on the growth ofDicorynia guianensis Amshoff seedlingsAnnals of Forest Science 57(7): 725–733[*INRA, Centre for Research Forestieres Nancy, F-54280 Champenoux, France]

Field assessment of composts produced by highly effective cellulolytic microor-ganismsBiology and Fertility of Soils 32(1): 35–40[*Natural Research Centre, Department of Agricultural Microbiology, Cairo, Egypt]

Interactions between a vesicular–arbuscular mycorrhizal fungus (Glomusintraradices) and Streptomyces coelicolor and their effects on sorghum plantsgrown in soil amended with chitin of brawn scalesBiology and Fertility of Soils 32(5): 401–409[*University of Mansoura, Faculty of Science, Department of Botany, Mansoura,Egypt]

The effect of long-term fertilization with organic or inorganic fertilizers onmycorrhiza mediated phosphorus uptake in subterranean cloverBiology and Fertility of Soils 32(5): 435–440[*CNRS, Centre Pedol Biology, 7 Rue ND des Pauvres, BP 5, F-54501 VandoeuvreNancy, France]

Evaluation of transcript level and biomass during mycelial development of thetruffle Tuber borchiiBiotechnology Letters 23(1): 17–20[*University of Urbino, Ist Chim Biol Giorgio Fornaini, Via Saffi 2, I-61029 Urbino,PU, Italy]

Arbuscular mycorrhizae respond to elevated atmospheric CO2 after long-termexposure: evidence from a CO2 spring in New Zealand supports the resourcebalance modelEcology Letters 3(6): 475–478[*University of Montana, Division of Biology Science, H S 104, Missoula, MT 59812,USA]

Bereau M, Barigah T S,Louisanna E, Garbaye J*. 2000

ElDin S M S B, Attia M*, andAboSedera S A. 2000

AbdelFattah G M* andMohamedin A H. 2000

Joner E J*. 2000

Zeppa S, Potenza L, ValloraniL, Guescini M, Ciarmela P,Stocchi V*. 2001

Rillig M C*, Hernandez G Y,and Newton P C D. 2000


Cloning and characterization of two repeated sequences in the symbiotic fun-gus Tuber melanosporum VittFEMS Microbiology Ecology 34(2): 139–146[*CNR, Inst Ric Miglioramento Genet Piante Foraggere, Via Madonna Alta 130, I-06128 Perugia, Italy]

Molecular approaches to arbuscular mycorrhiza functioningFungal Associations IX: 19–28[*Max Planck Inst Terr Mikrobiol, Karl Von Frisch Street, D-35043 Marburg, Ger-many]

Immunochemical characterization of mycorrhizal fungiFungal Associations IX: 29–43[*Tech University of Munchen Weihenstephan, Department of Botany, Alte Akad 12,D-85350 Freising, Germany]

At the interface between mycorrhizal fungi and plants: the structural organiza-tion of cell wall, plasma membrane and cytoskeletonFungal Associations IX: 45–61[*University of Turin, Dipartimento Biol Vegetale, CNR, Centre Studio MicolTerreno, Viale Mattioli 25, I-10125 Turin, Italy]

Lipids of mycorrhizaeFungal Associations IX: 63–93[*University of Littoral Cote Opale, LMPE, BP 699, F-62228 Calais, France]

Arbuscular mycorrhiza – a key component of sustainable plant–soil ecosystemsFungal Associations IX: 95–113[*University of Kent, Research School of Bioscience, Canterbury CT2 7NJ, Kent, Eng-land]

Exchange of carbohydrates between symbionts in ectomycorrhizaFungal Associations IX: 115–122[*University of Tubingen, Institute of Botany, Morgenstelle 1, D-72076 Tubingen,Germany]

Piriformospora indica: an axenically culturable mycorrhiza-like endosymbioticfungusFungal Associations IX: 125–150[*Jawaharlal Nehru University, School of Life Sciences, New Delhi 110 067, India]

Geosiphon pyriforme, an endocytosymbiosis between fungus and cyanobacteria,and its meaning as a model system for arbuscular mycorrhizal researchFungal Associations IX: 151–161[*Tech University of Darmstadt, Institute of Botany, Petersenstr 30, D-64287Darmstadt, Germany]

Mycorrhizal status in an orchard area in western Macedonia (Greece)Greece Agrochimica 44(3&4): 151–159[*NAGREF, Institute of Soil Science, Thermi Thessaloniki 57001]

Hydrolytic enzymes and ability of arbuscular mycorrhizal fungi to colonize rootsJournal of Experimental Botany 51(349): 1443–1448[*CSIC, Estac Expt Zaidin, Department of Microbiology Suelo & Sistemas Simbiot,Prof Albareda 1, Apdo 419, E-18008 Granada, Spain]



Paolocci F, Rubini A, RiccioniC, Granetti B, Arcioni S*.2000

Franken P* and Requena N.2001

Hahn A*, Wright S, and HockB. 2001

Bonfante P*. 2001

Sancholle M*, Dalpe Y, andGrandmouginFerjani A. 2001

Jeffries P* and Barea J M. 2001

Nehls U*, Wiese J, and HamppR. 2001

Varma A*, Singh A, Sudha,Sahay NS, Sharma J, Roy A,Kumari M, Rana D, ThakranS, Deka D, Bharti K, Herek T,Blechert O, Rexer KH, Kost G,Hahn A, Maier W, Walter M,Strack D, Kranner I . 2001

Schussler A* and Kluge M.2001

Karagiannidis N* and VelemisD. 2000

Garcia Garrido J M, Tribak M,Rejon Palomares A, Ocampo JA, Garcia Romera I*. 2000




The roles of auxins and cytokinins in mycorrhizal symbiosisJournal of Plant Growth Regulation 19(2): 144–154[*University of Western Australia, Faculty of Agriculture, Nedlands, WA 6907, Aus-tralia]

Evolution of mycorrhiza systemsNaturwissenschaften 87(11): 467–475[*University of Western Sydney, School of Science Food & Horticulture, MycorrhizaResearch Group, POB 10, Kingswood, NSW 2747, Australia]

Fungal enzymatic activity in fruitbodiesNova Hedwigia 71(3–4): 315–336[*University of Munich, Inst Systemat Bot, Sect Mycology, Menzinger Street 67, D-80638 Munich, Germany]

Distribution and molecular characterization of the root endophytePhialocephala fortinii along an environmental gradient in the boreal forest ofAlbertaMycological Research 104: 1213–1221[*University of Alberta, Department of Biology Science, Edmonton, AB T6G 2E9,Canada]

Polyamines in plant–microbe interactionsPhysiological and Molecular Plant Pathology 57(4): 137–146[*Scottish Agriculture College, Division of Plant & Crop Science, Department of PlantBiology, Auchincruive KA6 5HW, Ayr, Scotland]

Biodiversity: modelling angiosperms as networksPhytochemistry 55(6): 559–565[*FIOCRUZ, Inst Oswaldo Cruz, Dept Fisiol & Farmacodinam, BR-21045900 Rio DeJaneiro, Brazil]

Are ectomycorrhizal fungi associated with Alnus of importance for forest de-velopment in wet environment?Plant Biology 2(5): 505–511[*University of Nijmegen, Department of Aquat Ecology & Environment Biology,Toernooiveld 1, NL-6525 ED Nijmegen, The Netherlands]

Weaning performance of potato microplants following bacterization andmycorrhizationPotato Research 42(3&4): 521–527[*Natural University of Ireland University College Cork, Department of Plant Science,Cork, Ireland]

In vitro germination of nonphotosynthetic, myco-heterotrophic plants stimu-lated by fungi isolated from the adult plantsThe New Phytologist 148(2): 335–342[*University of California Berkeley, Department of Plant & Microbial Biology, 111Koshland Hall, Berkeley, CA 94720, USA]

Arbuscular mycorrhizal fungi might alleviate aluminium toxicity in bananaplantsThe New Phytologist 148(2): 343–352[*University of Catholique Louvain, BCCM, MUCL, Unite Microbiology, Pl CroixSud 3, B-1348 Louvain, Belgium]

Barker S J* and Tagu D. 2000

Cairney J W G*. 2000

Agerer R*, Schloter M, andHahn C. 2000

Addy H D, Hambleton S, andCurrah R S*. 2000

Walters D R*. 2000

Gottlieb O R and Borin M R DB. 2000

Baar J*, vanGroenendael J M,and Roelofs J G M. 2000

Duffy E M*, Hurley E M, andCassells A C. 1999

Bruns T D* and Read D J.2000

Rufyikiri G, Declerck S*,Dufey J E, and Delvaux B.2000

ISSN 0970-695X

Printed and published by Dr R K Pachauri on behalf of the Tata Energy Research Institute, Darbari Seth Block, Habitat Place, Lodhi Road,New Delhi – 110 003, and printed at Multiplexus (India), 94 B D Estate, Timarpur, Delhi – 110 054.

Editor Alok Adholeya Associate Editor Subrata Deb Assistant Editor Dr Vani Shanker

Forthcoming eventsConferences, congresses, seminars, symposiums, andworkshops

FAO/ECE/ILO Workshop on New Developments of Wood Harvesting with Cable SystemsR. Heinrich, Forest Harvesting, Trade and Marketing Branch, Forest Products Division,FAO, Viale delle Terme di Caracalla, 00100 Rome, Italy

Fax 39 06 5705 5137 • E-mail [email protected]

International Conference on Ex Situ and In Situ Conservation of CommercialTropical TreesMs Soetitah S. Soedojo, ITTO Project PD 16/ 96, Rev.4 (F), Faculty of Forestry, GadjahMada University, Bulaksumur, Yogyakarta 55281, Indonesia

Fax 62 274 902 220 • E-mail [email protected]

Second International Workshop on Edible Mycorrhizal MushroomsIan Hall E-mail [email protected] • Wang Yun E-mail [email protected]

Web site http://www.crop.cri.nz/whats_on/mushroom_conf

8th International Marine and Freshwater Mycology SymposiumSponsored by SouthValley University South Valley University, Hurghada, EgyptContact: Dr. H. H. El-Sharouny

E-mail [email protected] • Web site http://sites.netscape.net/botanyyoussufh/homepage

Third International Conference on MycorrhizasProf. Sally Smith, Department of Soil and Water, Waite Campus, The University of Ad-elaide, PMB 1, Glen Osmond, South Australia 5064

Fax +61 (08) 8303 6511 • Tel. +61 (08) 8303 7351 • E-mail [email protected] site http://www.waite.adelaide.edu.au/soil_science/3icom.html

Travelling Workshop on Linking the Complexity of Forest Canopies to Ecosystemsand Landscape FunctionMichael G. Ryan, USDA/FS Rocky Mountain Research Station, 240 West Prospect RD,Fort Collins, CO 80526-2098, USA

Fax 1 970 498 1027 • Tel. 1 970 498 1012 • E-mail [email protected]

Tree Biotechnology: the next millenniumDr Steven Strauss, Forestry Sciences Lab 020, Department of Forest Science, Oregon State,University, Corvallis, Oregon 97331-7501, USA

Fax 1 541 737 1393 • Tel. 1 541 737 6558 • E-mail [email protected] site http://www.cof.orst.edu/cof/extended/conferen/treebio/

The 2001 MSA meeting will be held with the American Phytopathological SocietyAbstracts from the 2000 meeting are still available on the MSA web site

5th International Flora Malesiana SymposiumDr Barry Conn, Royal Botanic Gardens, Sydney, and Mrs Macquaries Road, Sydney NSW2000, Australia

E-mail [email protected] • Web site http://plantnet.rbgsyd.gov.au/fm/fm.html

Dynamics of Forest Insect PopulationsDr Andrew Liebhold, USDA Forest Service, Northeastern Forest Experiment Station, For-estry Sciences Laboratory, 180 Canfield St., Morgantown West Virginia 26505, USA

Fax 1 304 285 1505 • Tel. 1 304 285 1609 • E-mail [email protected] site http://iufro.boku.ac.at/iufro/iufronet/d7/wu70307/aberdeen_firstannounce.htm

Improvement and Culture of EucalyptsIUFRO 2.08.03. Contact: Dr. Roberto Ipinza, Universidad Austral de Chile, POBox 1241, Valdivia, Chile

Fax 56 63 224 677 • Tel. 56 63 216 186 • E–mail [email protected]

MC 7 7th International Mycological CongressWeb site http://www.uio.no/conferences/imc7/

AustriaJune 2001

Yogyakarta, Indonesia11�13 June 2001

Christchurch, New Zealand3�6 July 2001

Hurghada, Egypt7�12 July 2001

Adelaide, Australia8�13 July 2001

Portland, Corvallis,USA11�19 July 2001

Skamania Lodge, Stevenson,Washington, USA22�27 July 2001

Salt Lake City, Utah1 September 2001

Sydney, Australia9�14 September 2001

Aberdeen, Scotland12�14 September 2001

Valdivia, ChileOctober 2001

Oslo, Norway11�17 August 2002

Regd No. 49170/89

About TERI The Bioresources and Biotechnology Division The...

Documents

Transcript of About TERI The Bioresources and Biotechnology Division The...