grain legumes no 49...no.37 genetic resources of grain legumes 3 r d q u a r t e r 2 0 0 2 no.34...

ISSUE No. 49 October 2007

AEPThe magazine of the European Association for Grain Legume Research

Le magazine de l’Association Européenne de recherche sur les Protéagineux

GRAIN LEGUMES

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Degradability of grain legumesin ruminants

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Environmental benefitsof grain legumes

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Genetic resourcesof grain legumes

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Highlights from the4th AEP Conference

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Back issuesBack issues

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Use of syntenyfor genetic progress

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The GRAIN LEGUMESIntegrated Project

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AEPThe magazine of th

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Le magazine de l’Association Européen

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Drought and saline stress in legumes

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Integrated activities in the EU Grain Legumes Integrated Project

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Vetches fromFeed to Food?

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Fababeans

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3 GRAIN LEGUMES No. 49 – October 2007

CONTENTS

Carte blanche4 Available markets should be supplied – we need to encourage

farmers to enlarge areas (P. Cuypers)

News5 Changes and challenges for AEP (The AEP Executive Committee)

5 Lisbon 2007 – Integrating legume biology for sustainableagriculture – a coming milestone

5 Insert: First GL-TTP workshop in April 2007

6 5th Phaseomics meeting in Varenna, Italy, May 2007 (F. Sparvoli, N. Terryn, W. Broughton)

6 Towards sustainability: eleventh Serbian forage crops symposium(A. Mikić)

Research7 Manipulating flowering time

(B. Wenden, C. Rameau I. Lejeune-Hénaut)

8 Insert: Digestibility of Hr winter peas (K. Crépon)

Special reportAbiotic stress in legumes9 Introduction10 Growth and functioning of legumes under drought: a whole

plant perspective (B. Davies)12 Plant traits and crop management to limit the effects of water stress

in Mediterranean agronomic situations (F. Lelièvre) 14 How genomics may help to understand tolerance to abiotic

stress in legumes? (V. Gruber, M. Crespi)16 Integrating molecular and genetic data in legumes (T. Huguet)17 Benefits of inoculation under abiotic stress in the Mediterranean

region (C. Vargas, M. E. Aouani)18 Abiotic stress breeding at ICARDA (M. Baum)19 Where we go from here – Concluding remarks of the abiotic stress

in legumes workshop(M. Crespi, A. Schneider, T. Huguet, M. E. Aouani)

Crops, uses & markets20 Dynamics and prospects for grain legumes in the European feed

market (A. Schneider, F. Pressenda, K. Crépon)22 Decision support system for efficient disease control

(T. Volk, J.-S. von Richthofen)24 A jump in price for French faba beans exported to Egypt

(J.-P. Lacampagne)

25 Events & new books

Around the world26 Pigeonpea development in China (S. Yang and colleagues)

The magazine of the European Association for Grain Legume Research

Le magazine de l’Association Européenne deRecherche sur les Légumineuses à graines

AEP12, avenue George V, 75008 Paris, France

Grain Legumes magazine is available on subscription to anyone interested

in grain legumes.

Subscription prices and ordering informationare available from the AEP secretariat or the

AEP web site(http://www.grainlegumes.com)

Publishing Director/Directeur de la publication:Álvaro RAMOS MONREAL

(Consejeria de Agricultura Ganadería, Spain)

Managing Editor/Rédacteur en chef: Anne SCHNEIDER (AEP, FR)

Email: [email protected]

Editorial Secretary/Secrétaire derédaction:

Jill CRAIG (AEP, UK)Email: [email protected]

Editorial Board/Comité de rédaction:Jill CRAIG (AEP, UK)

Gaëtan DUBOIS (UNIP, FR)Simon KIGHTLEY (NIAB, UK)

Pascal LETERME (Prairie Swine Centre,Saskatoon, Canada)

Wolfgang LINK (IPP, Göttingen, DE)Diego RUBIALES (CSIC, Córdoba, SP)

Anne SCHNEIDER (AEP, FR)

Overseas correspondents/Correspondants à l’étranger

Brian CLANCEY (Stat Publishing, USA)David MCNEIL (Victorian Department of

Primary Industries, AU)George HILL (Lincoln University, NZ)

Electronic prepress/InfographiePrinting/Impression:

Herald Graphics, Reading (UK)

Subscriptions/Abonnements:AEP Executive SecretariatEmail: [email protected]

Quarterly magazine/Revue trimestrielle

ISSN 245-4710

Commission paritaire n° 74 616

GG RR AA II NN LL EE GG UMEUME SS

4GRAIN LEGUMES No. 49 – October 2007

*UNIP President, Paris, France.([email protected])

Available markets should besupplied – we need to encouragefarmers to enlarge areas

Pierre Cuypers*

In the past three years, grain legume areas in France have decreasedsignificantly: 2007 areas are 50% less for peas and 30% lessfor faba beans compared with 2004 areas. In contrast to peas whoseyield also decreased, faba beans benefited from a quite valuable averageyield this 2007 harvest.

There has been a general trend in recent seasons for enhanced damageto arable crops caused by adverse climatic conditions during the cropcycle. However the decrease in grain legume area in France and inother EU countries is especially alarming for farming systems.

Indeed, as highlighted by Mr Christophe Terrain, President ofArvalis1 at a French press conference this summer, it is vital to managecropping decisions at the rotation level, and grain legumes bring aclear benefit in rotations. The Arvalis Acting Director, Gérard Moricepointed out that they enhance the yield of the following cereal by0.7 to 1 t/ha, and have costs €50 to €70 lower than a wheat oroil crop.

We need to respect the cycle of crops on one field, by avoiding thereturn of similar crops too soon, and maintaining the diversity ofcultivated species.

On the market side, UNIP was frustrated to see that there wasa low offer of faba beans and peas for the attractive export outletswhich have reached very high prices this commercial campaign!

Grain legumes compete indirectly with some other crops that benefitfrom a more positive image and from lucrative contracts, but we alsoidentified some logistical difficulties at collection, storage and marketlevels that are linked to minor volumes. That is why we believe indemonstrating actions towards farmers and farmers’ unions and ininnovative partnerships at regional levels to reinforce stability of thecrop chain from the grower to the market users.

In addition, we believe in collaboration with scientists to achievegenetic enhancement or agronomic solutions to overcome diseases andpests or soil and climatic variations, while maintaining high seed quality.

Grain legumes have benefits especially at agronomic andenvironmental levels, so we need to be creative to make themeconomically valuable! �1The technical institute for cereals and several other crops in France.

5 GRAIN LEGUMES No. 49 – October 2007

AEP NEWS

Changes and challenges for AEPSince autumn 2006, there have been some changes at AEP

headquarters.

A lighter structure for AEP in ParisOne of AEP’s main sponsors, UNIP, which hosts the AEP

headquarters, has experienced difficulties associated with the strongdecline in grain legume areas in recent years in France (as inother EU countries). UNIP has always supported AEP activitiesat financial and political levels and appreciates the successfulnetworking and dissemination efforts that have been carried outby AEP in the past years. However, with the new situation, UNIPcannot continue to support AEP without reductions in the activitiesand finances that go through the AEP office in Paris hosted byUNIP. The consequence is that the Paris headquarters of AEPnow have a lighter structure with only one half-time person andno more secretarial assistance.

In fact, UNIP proposes to take a greater part in AEP activitiesand decisions while ensuring the balance of AEP budgets. AEPhas always wished to involve more representatives from the industrysector. It is expected that this involvement will be followed bysimilar involvement of other market-related organisations oreconomic partners.

Therefore today the AEP network needs to find the best way(i) to operate with a lighter permanent structure for its day-to-day running as well as (ii) to meet the needs and demands of allthe different types of members within AEP.

New challenges for AEP after 15 years ofsuccessful development

Up to now AEP’s priority has been to establish an active networkin the legume scientific community, and AEP has been successfulin raising interest and funds for legume research at EU andinternational levels.

Now a new challenge is to develop additional activities of greaterinterest to economic players who face short and medium termproblems associated with the production and use of grain legumes;this means, identifying the technical issues to translate into questionsfor research for the short and medium terms, but also to preparelong term for the agriculture of the future.

At the same time, the activities of interest to scientists themselvesshould be maintained in AEP and the Executive Secretariat shouldfind a new way to function with smaller resources. This shouldbe discussed with any interested parties who wish to sustain suchan informal and neutral platform for collaboration, a joint voiceand the easiest dissemination of information. The joint activitieswill also be facilitated by the communication tools that havebeen developed by AEP in the recent years (web site, event cycles,GL magazine, etc).

Therefore facing the positive trends in R&D activities, thefuture AEP mandate will be an exciting and challenging periodfor legume enthusiasts!

Join AEP and apply for candidature to the Scientific Committeeto contribute to these challenges, facilitated by the previous effortsand success. �

Source: The AEP Executive Committee.

Lisbon-2007 – Integrating legume biology

for sustainable agriculture – a coming milestone for legumes

We are delighted to announce that our Lisbon-event to be heldfrom 12 to 16 November 2007 has received the patronageof Mr José Manuel Barroso, President of the European Commission.

Mr Timothy Hall, the Acting Director of Biotechnologies,Agriculture and Food of the European Commission Directorate-General for Research will officially open the conference and thePortuguese Minister of Agriculture and Rural Development, JaimeSilva, will give a welcome address on behalf of the PortugueseGovernment.

This event, the 6th European Conference on Grain Legumesfeatures the final dissemination event of the EU FP6 Grain LegumesIntegrated Project and promises to be a truly stimulating, scientificallyrewarding and sociably enjoyable experience for the 400 participants.

The programme is composed of plenary sessions, workshopsand poster displays, with presentations given by invited speakersand experts from a wide range of different fields, as well as plentyof opportunity for discussions.

This milestone will be the occasion to assess progress in grainlegume research and to prepare future collaborative work on legumes.

More up-to-date information about the programme is availableat: http://www.grainlegumes.com/index.php/aep/events/lisbon_2007/welcome_to_lisbon_2007 �

First GL-TTP workshop in April 2007The Grain Legumes Transfer Platform organised an exciting workshop entitled ‘TargetingScience to Real Needs” which attracted 63 participants from 17 countries (five continents) togather in Paris on 23–25 April 2007.

Reflecting the needs of grain legume breeders, the workshop focused on the exploitation ofgenetic resources, and on the concrete use and integration of molecular technologies inbreeding. The main themes addressed were genetic diversity, disease resistance, abiotic stresstolerance and quality traits.

As a follow-up to the event, GL-TTP is currently working with GL-TTP members to complete thetransfer of information and ready-to-use material presented at the workshop.

Finally, GL-TTP is building on the ideas that resulted from the brainstorming across professionalsectors to set up Research & Development and Technology Transfer projects in partnership withresearch scientists and plant breeders.

More information at http://www.gl-ttp.com


EURO AND WORLD NEWS

5th Phaseomics meeting in Varenna, Italy, May 2007

Towards sustainability: eleventh Serbian forage crops symposium

The 5th Phaseomics meeting was held in the beautiful settingof Villa Monasteria, Varenna, Lake Como, Italy. ‘Phaseomics’is a worldwide network of committed scientists that have ongoingresearch on beans. For more details visit www.phaseolus.net.

The highlights of the meeting were very diverse. There weresessions ranging from seed quality, to evolutionary genetics, genomics,genetic tools and resources, biotic and abiotic stresses, as well asthe interaction with beneficial organisms. Abstracts from themeeting, and interesting links can be found at www.phaseomicsv.net.

Of particular interest were talks from Matthew Blair (CIAT,Cali, Colombia) describing work on the use of wild accessionsto improve common bean varieties (1). This follows a trend setby work on rice (2) that uses wild accessions to improve crops.Despite plant breeders’ initial reluctance, the data shows clearlythat this is a process that also ‘yields’ in grain legumes!

Other interesting talks covered the food and feed qualities ofbean (Francesca Sparvoli and Bruno Campion, Milano, Italy).They have developed and are now evaluating mutants with lowphytic acid contents as well as beans that are lectin-free. Bothcould help improve the digestibility of beans. Nutritional workis in progress to verify this hypothesis.

As for genomics tools, a TILLING population in the varietyBAT93 is being set up by collaboration between three teams(CIAT, University Geneva and USDA/ARS/TARS Puerto Rico).Part of the bean genome will be sequenced and different groupshave contributed to the growing number of bean ESTs in Genbank.

In general the meeting was very successful, and the feelingwas that the future for beans, as the most important food legume,looks very positive. �

Source: Francesca Sparvoli ([email protected]), Nancy Terryn([email protected]) and William Broughton([email protected])

(1) Blair, M. W. et al. (2006). Theoretical Applied Genetics 112, 1149–1163.

(2) McCouch, S. R. et al (2007). Euphytica 154 (3), 317–339.

The eleventh symposium on forage crops of the Republic ofSerbia was held from 30 May to 1 June 2007 at the Facultyof Agriculture in Novi Sad under the auspices of the SerbianMinistries of Science and of Agriculture, Forestry and WaterManagement. The symposium was organised by the Forage CropsSociety of Serbia, the Institute of Field and Vegetable Crops(IFVCNS) in Novi Sad and the Faculty of Agriculture of theUniversity of Novi Sad (FANS).

With the title ‘Systems of sustainable production and utilisationof forage crops’, the meeting brought together more than 100researchers, with about 70 from Serbia and more than 30 from14 other countries. The three main sessions focused on (1) breeding,seed production and genetic resources, (2) field forage cropsproduction and utilisation and (3) grassland production.

Half the papers were on legumesAbout half of the one hundred papers presented were related

to legumes, confirming the essential role that these crops, togetherwith grasses and other forage crops, play in modern systems offeed production. Some papers focused on single species, and dealtmostly with lucerne, followed by vetches and red clover, whileothers focused on breeding and agronomy.

Following plenary talks on recent achievements and the futuredirection of breeding perennial and annual legumes in Serbia(Dragan -Dukić , FANS, Serbia and Vojislav Mihailović, IFVCNS,

Serbia) and also a review of the current status and prospects forannual forage and grain legumes in contemporary Serbian agriculture(Branko Ćupina, FANS, Serbia), other authors from Serbia presentednumerous interesting contributions on the diversity of legumespecies and their utilisation. Among the highlights of the first twosessions were talks by foreign delegates: the genetics of agronomiccharacteristics in pea (Noel Ellis, JIC, UK), improving forageand seed yield in temperate legumes (Athole Marshall, IGER,UK), common vetch breeding (Rade Matić, SARDI, Australia),breeding sunn hemp (Crotalaria juncea) and sericea lespedeza(Lespedeza cuneata) (Jorge Mosjidis, Auburn University, USA),various cultivar evaluations (Fred Eickmeyer, Saatzucht Steinach,Germany and Piotr Stypiński, Warsaw Agricultural University,Poland), forage potential of a grain legume collection (MargaritaVishnyakova, VIR, Russia), lucerne management in irrigated areas(Jaume Lloveras, University of Lleida, Spain), the tripartite symbioticsystem in legumes (Alexey Borisov, ARRIRAM, Russia), the roleof legumes in sward management (Milus̆e Svobodová, CzechUniversity of Agriculture) and red clover and other crops asalternatives to grasses for ensilage (Pádraig O’Kiely (Teagasc, Ireland).

All the symposium papers were published as Volume 44 of thejournal, A Periodical of Scientific Research on Field and Vegetable Crops,issued by the Institute of Field and Vegetable Crops and freelyavailable on a CD in pdf. format from Aleksandar Mikić. �

Source: Aleksandar Mikić ([email protected]).

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GRAIN LEGUMES No. 49 – October 20077

RESEARCH

*SGAP INRA, Versailles, France.([email protected],[email protected])

**UMR INRA/USTL SADV, Estrées-Mons,Péronne, France. ([email protected])

Manipulating flowering timeChanger la date de floraison du poisby Bénédicte WENDEN*, Catherine RAMEAU* and Isabelle LEJEUNE-HÉNAUT**

Pea, the most representative Europeangrain legume has limited developmentprospects mainly due to unstable yieldswithin years or sites. Stabilising andimproving the productivity of the dry peacrop could be achieved through the releaseof winter cultivars, as has been done, forexample, for wheat and canola.

Towards an ideal winter pea Escape from drought and heat stresses in

the spring and through the earlier floweringperiod, together with a longer developmentcycle leading to higher global biomassproduction, would make the productivityof such winter grown cultivars more reliable(Figure 1).

For an autumn-sown pea however, it isessential to overcome frost which is anotherimportant abiotic stress. For the last 30years, winter dry pea cultivars have beenbred progressively for better frost tolerance,but until now they have never reachedthe level of tolerance expressed by someforage lines, like Champagne or AustrianWinter. Given that frost sensitivity risesafter floral initiation, previous observationshave shown that such frost tolerant peas areable to escape the main winter freezingperiods by delaying their floral initiationunder short days (3).

For the winter pea crop, themanipulation of the flowering genes appearsto be a promising way to control the keydevelopmental stages which are the datesof floral initiation and flowering. The idealwinter pea should initiate its flowerprimordia late enough to avoid winter frostsand should flower early enough to escapedrought and heat stresses in late spring.

Regulating flowering time Since the early 1950s, pea has been used

as a model species for the study of thegenetic control of flowering time throughthe physiological characterisation of mutantsobserved in different environmentalconditions (long days, short days) and theuse of grafting experiments. These studiesresulted in a classical model which iscurrently being reconciled with themolecular genetic models developed inArabidopsis by J. Weller’s group at theUniversity of Tasmania. The fact that mostof the flowering genes identified inArabidopsis are present in legume genomesequence databases (2) suggests that similarmechanisms are likely to occur betweenpea and Arabidopsis, and that already wehave the tools to understand and to controlflowering time in pea. In particular, theresponse to photoperiod in pea could, asfor Arabidopsis, also involve the circadianclock and the precise cyclic regulation ofphotoperiodic genes (CONSTANS) (5).

The winter pea breeding strategydeveloped at INRA relies partly on themanipulation of two major genescontrolling the natural variation forflowering time, namely LF and HR. Thesegenes are promising targets because theyare known to control, respectively, theintrinsic earliness for the beginning offlowering and the strength of the response

to the photoperiod. More precisely, thedominant allele HR is known to delay thefloral initiation of autumn-sown peas untila longer daylength is reached (approximately13 h 30 min) in the following spring. Thishelps to escape the main winter freezingperiods. The knowledge of the sequenceof these genes will allow the developmentof molecular markers suitable for buildingoptimal associations of the HR and LFalleles. A candidate gene approach hasalready led to the cloning of the LF geneas the homologue of the TFL1 Arabidopsisgene (1), and molecular markers are beingdeveloped to follow the different allelesin crosses. For HR the same approach isbeing used, thanks to the microsyntenywith Medicago truncatula, in order to clonethe gene and/or develop molecular markersto backcross the high photoperiod responsedominant HR allele in agronomic geneticbackgrounds.

Developing integrative toolsTo understand the biology of complex

systems it is necessary to integrate atdifferent scales the existing models,experimental data, and main hypothesesinto new models that are able to describe,explain and predict biological phenomena.A PhD research project is ongoing at INRAVersailles to develop an integrative toolcombining molecular, physiological and

Spring pea

Traditional winter pea

Winter pea ideotype

Autumnrains

Wintercold stress

Sowing Floralinitiation

Yield maturity

Heat and droughtstresses

HR :Photoperiod

sensitivity

LF :Endogenous

precocity

Figure 1. The winter pea strategy: manipulating the dates of floral initiation and flowering in order to escapewinter frosts as well as drought and heat stresses in late spring.

RESEARCH


ecophysiological data into a computer modelof flowering time in pea. This model willintegrate the hypotheses that best representour current understanding of the geneticand physiological regulation of floweringtime from the genetic to the crop level.

Presently, two categories of floweringmodels can be described for pea. First, anagroecophysiological model has beenestablished, primarily for two pea varieties,in which the time of flowering has beenbroken down into two componentvariables: the leaf appearance rate and thenode of the first open flower (4). However,the response of flowering to theenvironment also depends on genotype andtherefore extrapolation of these results toother environmental conditions orgenotypes is limited. Secondly, geneticmodels of the control of flowering in peahave been based on the analyses of floweringmutants (see above) describing theinteractions of signals that lead to theproduction of a flowering signal and thus

to floral initiation (6), but these geneticmodels have no prediction capacity.

The aim of the PhD project is to gatherall the sources of knowledge in the formof mathematical equations that will togetherallow the simulation of as many differentenvironmental and genetic conditions aspossible. In particular, this work shouldenable the effects of the different genes,and their interaction with environmentalconditions, on flowering time to bequantified. The modelling approach is basedon two sets of data (Figure 2): first thegenotype of the key flowering genes (LF,HR, SN, DNE, GI) and secondenvironmental data (photoperiod and meantemperature). For this model, we assume,as it was shown in Arabidopsis, that theproduction of the flowering signal isquantitatively controlled by an integratednetwork of pathways. The response offlowering to photoperiod is modelledthrough the effect of cycling genescontrolled by the circadian clock and the

coincidence between their level ofexpression and the daylength. Theflowering signal accumulates over time untilit reaches a flowering threshold, determinedby the LF allele, and then triggers floralinitiation at the apex. The main outputs ofthis integrative model are the node andtime of floral initiation as well as the nodeand time of flowering. These variables couldin turn be integrated as a prediction offlowering into a crop growth model.

How can this model be used?Knowledge of the genetic and

physiological basis of the interactionbetween flowering genes and theenvironment is essential to the selectionof varieties closely adapted to a specificenvironment. This work will lead first toa better comprehension of the effects ofthe different flowering genes in pea, andtheir interaction with the environment.

Then, with a better understanding andprediction of flowering time, new ‘virtual’pea genotypes could be constructed andtheir flowering phenotypes predicted underdifferent environmental conditions. Bytesting different LF/HR allele associations,under virtual winter conditions, we will beable to predict the best winter pea floweringgenotype for a given environment and toprovide precise targets to the breeders. �

(1) Foucher, F. et al. (2003). Plant Cell 15,2742–2754.

(2) Hecht, V. et al. (2005). Plant Physiol 137,1420–1434.

(3) Lejeune-Henaut, I. et al. (1999). Euphytica109 (3), 201–211.

(4) Truong, H. H. and Duthion, C. (1993). AnnBot. 72(2), 133–142.

(5) Weller, J. (2005). Flowering Newsletter 40,39–42

(6) Weller, J. L. et al. (1997). Trends in PlantScience 2 (11), 412–418.

Data Integrative tools Predictions

GenotypeFlowering keygenes

•LF•HR

Environment

Photoperiod

Temperature

ModellingGenetic network

Circadianclock

Otherpathways

SignalsCycling genes

Production of a flowering signal

Flowering threshold (LF)

Time or Node(developmental rate)

Node offloral

initiation

Floralinitiation

time

Floweringnode

Floweringtime

Figure 2. An integrative model to predict the flowering time in pea.

Insert

Digestibility of Hr winter peasPea breeders have started to breed varieties sensitive to photoperiod(length of the daylight) since they show an advantageous longerperiod of frost tolerance. This photoperiod sensitivity is controlledby the Hr gene.

In parallel, the nutritional value of these types of peas for pigshas been tested by UNIP and Arvalis Institut du Végétal in France,by comparing one classical spring pea (SP) to both one linesensitive to the photoperiod (HR1) and one line with a better

agronomic value (HR2) issued from the cross between HR1 and aspring variety.

The two winter type varieties (HR1 and HR2) have a lower trypsininhibitor activity compared with the one of spring type. The aminoacid profiles of HR1 and HR2 are similar and slightly different fromthe one of SP but are very close to the usual value for peas.

The protein digestibility is 82% in HR2, slightly higher than for HR1 (80%) and SP (78%) but without significant difference.

The amino acid digestibility is high and similar in the three types.

Therefore the genetic character which modifies the timing ofthe floral initiation does not modify the chemical compositionor digestibility of the proteins and amino acids in pigs.

Source: Katell Crépon, UNIP, [email protected]


Legumes are sensitive to abiotic stresses such as waterdeficit and soil salinity. Tolerance and susceptibility to thesestresses are very complex traits and plants show complexcell signalling pathways activating metabolic functions anddevelopmental switches to cope with environmental stresses.

Hence, international experts working on the analysis ofplant tolerance were gathered at a scientific workshop ‘Abioticstresses in legumes’ (Tunis, 22–24 March 2007, supported byGLIP1). Basic scientists, molecular biologists, genomicists,physiologists and agronomists involved in several ongoingEU projects were present together with representatives ofinternational agricultural organisations. Legume experts andtheir colleagues working on non-legume plants came togetherso as to learn from a variety of stress response models:agronomic, eco-physiological and genomic models. In thelatter case, the well known Arabidopsis model has allowedtremendous progress in the genomic analysis of abiotic stresseseven though several physiological characteristics of legumesare not found in this plant. More recently Medicago truncatula,an annual, diploid and autogamous legume, has been adoptedas a model legume in various European and USA laboratoriesand diverse genetic, genomic and reverse genetic tools arebeing developed for the analysis of physiological responsesat genomic level.

This special report2 brings together some highlights of thisworkshop and presents current perspectives of the way plantscope with abiotic stresses with particular emphasis on theimplications for grain legume crops. The data discussed dealwith whole-plant approaches and crop management as well asgenetic and genomic analysis of abiotic stress responses. �1the EU Grain Legumes Integrated Project www.eugrainlegumes.org 2a previous dossier dealt with the same topic: Special report (2005)Drought and saline stress in legumes, Grain Legumes 42, 13–22.

Les légumineuses sont sensibles aux stresses abiotiques commele déficit hydrique et la salinité du sol. La tolérance et lasusceptibilité à ce type de contraintes sont des caractèrescomplexes et les plantes présentent des processus d’échangescellulaires élaborées, activant des fonctions métaboliques et deschangements de développement pour gérer les stressesenvironnementaux.

Il a été décidé de rassembler plusieurs experts internationauxtravaillant sur l’analyse de la tolérance végétale à un séminairescientifique ‘Les stress abiotiques et les légumineuses’ (Tunis,22–24 mars 2007, co-organisé par GLIP1), avec des scientifiquesplus fondamentaux, des génomiciens, des physiologistes etdes agronomes. Les experts légumineuses et leurs collèguesspécialistes des plantes modèles ont partagé leurs différentsschémas d’analyse de la réponse végétale au stress :agronomiques, éco-physiologiques ou génomiques. La plantemodèle Arabidopsis a notamment permis des avancéesconsidérables sur la génomique de la réponse au stress mêmesi cette plante n’a pas certaines fonctionnalités des légumineuses.Plus récemment, Medicago truncatula, une légumineuse annuelle,diploïde et autogame, a été adoptée comme modèle deslégumineuses par différents laboratoires européens et américains,et des outils génétiques et de génomie fonctionnelle ont étédéveloppés pour l’analyse des réponses physiologiques.

Le dossier de ce numéro2 rapporte quelques informations saillantesdu séminaire de Tunis, analysant les différentes façons dont lesplantes gèrent les stress abiotiques, en insistant sur les conséquencessur les cultures de légumineuses. Sont traités à la fois les approchesintégrant la gestion de la culture, la compréhension de laphysiologie de la plante dans son ensemble et les analysesgénomiques des réponses de la plante au stress. �1le projet intégré européen sur les légumineuses à graines www.eugrainlegumes.org 2un dossier précédent traitait aussi de ce sujet: Special report (2005) Drought andsaline stress in legumes, Grain Legumes 42, 13–22.

Salt effect on the growth of Medicago truncatula F83005.5 line. Left: no salt, right: 45 mM NaCl. Abiotic stress in field trial of faba beans at El-Sharkiain Egypt.

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SPECIAL REPORT

Abiotic stress in legumesLes stress abiotiques et les légumineuses

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Abiotic stress in legumesLes stress abiotiques et les légumineuses


SPEC IAL REPO

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ABIOTIC STRESS

fixation, vegetative growth and developmentand aspects of reproductive developmentare most sensitive to soil drying.

A high leaf area index is vital to maximisethe interception of radiation forphotosynthate production. Restrictions incanopy development under drought, andlimitation in vegetative growth before a plantcommunity has covered the ground can bevery damaging for carbon (C) accumulationand yielding. Furthermore, covering thesoil early in the growing season can restrictsoil water loss significantly and therebyenhance the transpiration (T) componentof evapotranspiration (ET), resulting inmore productive use of water (16). Onlywater taken up by the plant can contributeto C gain (W in Equation 1 below).

TurgorCellular water status (turgor) provides

the driving force for organ growth andusually leaf growth becomes restricted asthe soil dries because leaf turgor falls belowa threshold value for leaf expansion. Somegenotypes accumulate solute in cells as shootwater potential falls, thereby maintainingturgor and the potential for growth.However, the maintenance of leaf turgordoes not usually result in maintenance ofshoot growth as the soil dries. A goodcorrelation between restricted shoot growthand high solute accumulation in shootsindicates that some variable other than cellturgor can restrict cell growth under drought.In this situation solutes accumulate as a resultof restricted leaf growth, rather thanproviding the driving force for cell expansionas the soil dries (7). Solute accumulationcan prolong the period before cells die andin some dryland legumes, postponing celldeath by this method is adaptive (5).

All crop plant species are highlysensitive to soil drying, withreductions in productivityoccurring before the amount of wateravailable to the plant has changedsignificantly. Sensing involves both reduceduptake of water from the soil and/or themodified production and transport of a rangeof chemical signals in roots in contact withdrying soil. Both of these responses canprovide information to the shoots on soilwater availability. These changes in chemicaland hydraulic signalling allow plants toregulate growth and development as afunction of resource availability to the roots.

Plant breeders believe that characteristicsimportant for yield in resource-poorenvironments have more to do with growthunder favourable conditions than withresistance to stress per se. As there is littlegrowth by plants once stress develops, overallproductivity is best improved by maximisinggrowth in favourable times (10). Processesthat help the plant survive severe cellulardehydration may be important in perennialcrops but to sustain yields of annuals weneed to understand the regulation of growthand development under moderate droughtstress, rather than focussing on survival andtolerance of low plant water content.

Drought regulates a varietyof plant processes

The sensitivity of different aspects of theplant’s physiology and biochemistry todrought can vary substantially; lesions inmost processes will contribute to reducedyields and/or crop quality. Nitrogen (N)

There is often some root growth in dryingsoil, even when shoot growth is entirelyrestricted. As a result the plant root:shootratio is increased. The development of waterand nutrient depletion zones in soilsurrounding slowly growing roots(particularly when roots clump in compactedsoil) restricts resource uptake from dryingsoil. In these circumstances sustained rootgrowth sustains water and nutrient uptake.

Selection for solute regulation capacityin roots may be a way of developing cropsfor dryland regions (11) and many haveshown the positive impact of root turgormaintenance on growth (3). Sustained rootgrowth in drying soil may also requireaccumulation of the plant hormone abscisicacid (ABA) which prevents run-awayethylene production at low water potentials

Growth and functioning of legumes under drought: a whole plant perspectiveCroissance et fonctionnement des légumineuses sous déficithydrique : une approche à l’échelle de la planteby Bill DAVIES*

*Lancaster Environment Centre, LancasterUniversity, UK. ([email protected])

What is water stress?Successful growth and development of landplants requires maintenance of a favourablecellular water status in even the mostchallenging of climates. The dry airsurrounding the above-ground parts of theplant promotes rates of water loss(transpiration) from the shoots that potentiallycan be very high. Plants must protectthemselves against excessive water loss (usingwater-proofing of aerial plant surfaces andstomatal pores of variable aperture). This willhelp avoid the damaging effects of stresswhich will inevitably limit all aspects of growthand development and ultimately result in celldeath. These stresses may be low cell watercontents (low water potential) and also thebuild-up of growth-inhibiting chemicals.Sustained water uptake from soil and transportof this water to leaves is also required toreplace water lost via transpiration andavoid the development of damaging deficits.


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(15). In many circumstances ethylene is apowerful growth inhibitor (13) and, as thesoil dries, increased soil strength may alsolimit shoot growth through up-regulationof ethylene synthesis. Ethylene accumulationcan also restrict nodulation significantly.Low ethylene transgenics show reducedleaf growth sensitivity to compacted soiland to soil at low water potential (14).

Three key cropping variablesAn analysis by Passioura (9) highlights

three key cropping variables that willcontribute to sustained crop yield (Y) underdrought:

Y = WUE x W x HI (Equation 1)

WUE = plant biomass produced per unitof water used, W = water available, and HI= harvest index (proportion of biomassproduced as grain). We have seen abovehow W can be manipulated to sustain Y.WUE can be enhanced, for example, byselecting for or manipulating particularaspects of photsynthesis or stomatalbehaviour. Crop yield can also be sustainedby ensuring that water is available at keydevelopmental periods so that a greaterproportion of crop biomass is yield (HI).

Recent work shows the importance ofsustaining C supply to developing maizegrains immediately after anthesis. These Callocation processes are extremely sensitiveto water availability and if water relationsare adverse during this critical period ofcrop development then grain yield fails (1),in spite of favourable water relations anddevelopment of the vegetative crop.

N fixation and droughtSoil drying can reduce nodule mass

substantially, having a negative impact on Naccumulation through fixation. Interestingly,Rhizobia are quite resistant to soil dryingand their survival is probably not a limitingstep for N fixation under drought, eventhough symbiosis establishment is extremelysensitive to drought. Despite the apparentrobustness of Rhizobia in water-limitedenvironments, data indicating superiorperformance of some, often indigenous,strains in dryland environments are limited.

In many legumes, N fixation per unitmass of nodules is highly sensitive to drought,often more sensitive than other plantphysiological and developmental processes(12). Carbon and oxygen limitation in thenodules and/or feedback regulation of N

fixation by N accumulation are possibleexplanations for this sensitivity, linked directlyto changes in the water status of the plant.However, nodules obtain most of their waterfrom the phloem rather than the xylem.Variation in phloem flow can be highlysensitive to small changes in plant water statusand can impact severely on nodule activity.

Variation in phloem and xylemfunctioning could also affect chemicalregulation of nodule activity. For example,if an increase in the concentration of phloemcontents occurs at reduced fluxes, assimilatesdelivery to nodules may be sustained.However, export of N products from nodulesvia the xylem may be decreased by stress,resulting in N accumulation and N feedbackinhibition of nodule activity. Suppressionof N fixation of this kind can result fromlocal signalling (8) or a more systemic signalimported by the phloem (12).

Manipulation of chemicalsignalling

There is significant genetic variationbetween cultivars in N-fixation sensitivity tosoil drying, but there is still some uncertaintyover the basis of this variation. Until thereis some agreement over the mechanismby which soil drying reduces N-fixationactivity, the opportunities for manipulatingthis activity will remain restricted, and wewill not be able to benefit from thedevelopment of novel biotechnologicaltechniques for plant improvement.

Increased understanding of chemicalsignalling mechanisms in drought-affectedplants raises the possibility that artificialmanipulation of signal synthesis andaccumulation may enhance plantperformance when water is restricted. Thismay be achieved via plant improvement orvia a change in management practice, forexample, via deficit irrigation. One of thesetechniques, partial root drying, has beendesigned so that some plant roots are alwaysin contact with drying soil and will thereforecontinue to produce chemical signals. Usingdeficit irrigation, signal production can beenhanced or decreased at critical periodsof plant development resulting in restrictedshoot growth and water use with very littlelimitation in reproductive yield (increasedHI – Equation 1) (4). Other manipulations,such as addition of specific bacteria to therhizosphere (2) or fertiliser treatments willaugment or restrict other chemical signals

in plants. These could, for example, ensurerapid canopy development in drying soil(16) or ultimately promote plant senescencethereby enhancing partitioning of C andN to increase HI of grain crops (17).

Opportunities for plantbiologists

A range of plant stress-sensing mechanismshas evolved so that, during mild soil drying,plants start to shut down growth andfunctioning to save scarce water for criticalphases of their development, even whenthere is still water available in the soil. Thissuggests that there is some scope for plantbiologists to over-ride these defensiveresponses with the benefit of sustained plantdevelopment and yielding as the soil dries.Further elucidation of the drought-sensingsystem to increase the efficiency of wateruse in agriculture (‘more crop per drop’)is most relevant as rainfall patterns aredisrupted by climatic change. �

(1) Boyle, M. G. et al. (1991). Crop Science 31,1246–1252.(2) Belimov, A. A. et al. (2007). Journal ofExperimental Botany 58, 1485–1495.(3) Davies, W. J. (2007). In: Advances inmolecular breeding towards salinity and droughttolerance, 55–72 (Eds M. A. Jenks et al.).Springer, Heidelberg, Germany.(4) Davies, W. J. et al. (2002). New Phytologist153, 449.(5) Flower, D. J. and Ludlow, M. M. (1986).Plant Cell and Environment 9, 33–40.(6) Kirda, C. et al. (1989). Plant and Soil 120,49–55.(7) Kuang, J. B. et al. (1990). Journal ofExperimental Botany 41, 217–221.(8) Marino, D. et al. (2007). Plant Physiology143, 1968–1974(9) Passioura, J. B. (1977). Journal of theAustralian Institute of Agricultural Science 43,117–121.(10) Richards, R. J. (1993). In: Plant responsesto cellular dehydration, 211–223. (Eds T. J.Close and E. A. Bray). American Society ofPlant Physiologists, Rockville, USA.(11) Serraj, R. and Sinclair, T. R. (2002). PlantCell and Environment 25, 333–334.(12) Serraj, R. et al. (1999). Journal ofExperimental Botany 50, 143–155.(13) Sharp, R. E. (2002). Plant Cell andEnvironment 25, 211–222. (14) Sobeih, W. et al. (2004). Journal ofExperimental Botany 55, 2353–2364.(15) Spollen, W. G. et al. (1993). In: Waterdeficits: plant responses from cell to community,37-52. (Eds J. A. C. Smith and H. Griffiths)BIOS Scientific, Oxford, UK.(16) Turner, N. C. (2004). Journal ofExperimental Botany 55, 2413–2425.(17) Yang, J. (2001). Agronomy Journal 93,196–206.


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Plant traits and crop managment to limit the effectsof water stress in Mediterranean agronomicsituationsCaractères des plantes et conduite des cultures pour limiterl’impact de la sécheresse en agriculture méditerranéenneby François LELIÈVRE*

*INRA, UMR SYSTEM, Montpellier, France.([email protected])

Water deficit is the main constraintfor agricultural production inMediterranean environments.Genetic improvement and agronomicoptimisation of harvested yield, HY(g DM/m2) depend on three termsaccording to the relation (Equation 1)proposed by Passioura (3, 4):

Equation 1: HY = T x WUE x HI (with DMt = T x WUE)

where T (g H2O/g DM per m2) is the croptranspiration, WUE is average water useefficiency (g DM/kg H2O), HI is the harvestindex, and DMt (g DM/m2) is the totaldry matter elaborated. If E (g H2O/m2) isthe soil evaporation, ET = E+T is the actualevapo-transpiration of the crop during itscycle. WUE is often related to aerial drymatter, DMa, instead of DMt (aerial +underground). All these variables are sumsor means for the whole cycle of the crop,representing several months or even yearswhen pluri-annual yields are considered.The improvement of each term T, WUEand HI has limits which are discussed.

Increasing and stabilisingharvest index

In annual grain crops in all environments,yields have progressed in the last half centurythrough continuous genetic HI increase(from 0.1–0.2 to around 0.5 in manyspecies). In rainfed semi-arid environments,subsequent improvement of HI stability has

been obtained in winter cereals by plantbreeding through: (i) genetic control ofvegetative growth and stem length; (ii)drought resistance of pollination-fertilisationphase; (iii) drought escape of the grain fillingperiod through earliness and deep rooting;(iv) increased resistance of late photosynthesisand late translocation during grain filling;(v) improved diseases resistance. Much lesssignificant are the results obtained in HIstability of grain legumes, but progress shouldbe obtained by following the same objectives.

Optimising crop transpirationCrop transpiration (T) can be increased

through plant cycle duration (earliness)within limits in one year. In temperate areas,annual rainfall (Pa) generally exceeds annualpotential evapo-transpiration (ETpa); actualtranspiration (Ta) is limited by ETpa(Ta≤ETpa≤Pa). In Mediterranean areas ETpaexceeds Pa which is the upper limit of Ta(Ta



where the non-growing period is winter. Recent work on wheat in semi-arid

conditions has demonstrated that earlyautumn sowing combined with genotypeswith high leaf and root extension rates at low temperatures (autumn, winter, early spring) are very efficient to increaseTa if the balance between reproductive and vegetative growth is controlled bygenetic factors (6). A similar approach is valuable for winter grain legumes in the Mediterranean.

Optimising water useefficiency

Water use efficiency (WUE) is theeffectiveness of a crop in H2O/CO2exchange, the value representing a verylong and complicated process ofdevelopment and growth in real conditions(1). Values of 3–5 g DM/kg H2O representhigh utilisation of water by agriculture,from which approximately a half can beallocated to seeds in grain crops. Such levelsare not possible everywhere at any time,because WUE follows biophysical laws: itdecreases under high temperatures and highclimatic demands (2–3 times lower in hotcompared with cool seasons/climates). Inhot periods or climates, it is higher in C4than in C3 plants. High nutrient availabilityincreases WUE, especially nitrogen forcereals and grasses and phosphorus for

legumes. In North Africa, WUE ofirrigated alfalfa is 3.0–4.0 g DM/kg H2Oin spring but falls to 0.5–1.0 in summer.In annual legumes, adaptation to earlyautumn sowings with efficient nitrogenfixation and growth at low temperaturesis important. Conventional selection forDM yield in rainfed field plots selectsefficiently for WUE because high yieldingmaterial of similar earliness useapproximately the same quantity of water(2). However, conventional breeding forDMt and WUE increase is slow becauseof low variability in high yielding materialin a species. Molecular engineering shouldprovide opportunities to modify H2O/CO2exchange (1).

The case of a perennialforage crop

Harvested yield for one year (HYa b =Ta x WUEa) is generally the sum of severalcuts (growth cycles at different developmentperiods having different daily Tj and WUEj.For pluri-annual forage crops like alfalfa,yield is the sum during n years: HYn = Sn(HYa).

In temperate areas, forage yield HYa ismaximised when Ta tends to ETPa (dailyTj = ETPj throughout the year). It requires:(i) high plant density; (ii) early start ofgrowth in spring and late end in autumn;(iii) drought resistance in summer. Plants

must have the ability to maintain optimalor significant T, growth and yield duringmoderate dry periods that do not exceedtwo months with cumulated climaticdeficits (ETp-P) not lower than –250 mm.It is obtained through depletion of deepsoil water reserves, mainly conditionedby soil and root depth.

In rainfed Mediterranean areas, adaptedplants must alternate: (i) a productivestrategy with high T and high WUE duringthe cool rainy season (5–9 months, P:300–700 mm, ETP: 300–600 mm); (ii) aconservative strategy (survival) withoutgrowth, with partial senescence and verylow T, during the long hot dry season (3–7months, P: 0–100 mm, ETP: 400–1000mm). During this period, surviving organslose water which is compensated for byextraction from the soil to maintainhydration above the lethal level for as longas possible. This conservative transpiration(Ts), not used for direct production, isthe ‘water cost’ of summer survival andperenniality, in addition to soil evaporation.Adaptation in perennials depends on acombination of plant traits like deep rooting,different levels of summer dormancy, andrelative desiccation tolerance of survivingorgans (7, 8). Interest and limits of perennialvs. annual (reseeded or self-reseeding) foragelegumes and grasses must be evaluated atthe farming systems level. In this area, thegap from plant physiology and moleculargenetics studies to the availability ofinnovative cultivars as commercial seedmust be reduced. �

(1) Bacon, M. A. (Editor) (2004). Water useefficiency in plant biology. Blackwell PublishingCRC Press, Oxford, UK. 327 p.

(2) Lelièvre, F. (2006). In: Proc. of Eucarpiaconf., fodder crops and amenity grasses section,Peruggia, Italy, 3–7 Sept 2006, 279–283 (Eds D.Rossellini and F. Veronesi). Univ. di PerugggiaPubl., Peruggia, Italy.

(3) Passioura, J. B. (1977). J. Austr. Inst. ofAgric. Sc. 43, 117–121.

(4) Passioura, J. B. (2002). Functionnal PlantBiology 29, 537–546.

(5) Passioura, J. B. (2004). In:Water useefficiency in plant biology, 302–321 (Ed. M. A.Bacon). Blackwell Publishing, Oxford, UK.

(6) Richards, R. A. et al. (2002). Crop Science42, 111–121.

(7) Volaire, F. et al. (2005). Annals of Botany 95,981–990.

(8) Volaire, F. and Norton, M. (2006). Annals ofBotany 98, 927–933.

Trial to intercrop vineyards with perennial legumes and grasses on shallow soils in French Catalogne (southern France).

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How genomics may help to understand tolerance toabiotic stress in legumes? Comment la génomique contribue t-elle à comprendre latolérance aux stress abiotiques chez les légumineuses?by Véronique GRUBER* and Martin CRESPI*

soil generate a low water potential zonemaking it increasingly difficult for the plantroots to acquire both water and nutrients.Both hyperionic and hyperosmotic stressinduce drastic metabolic changes that canlead to plant death. In legumes, salt stressalso inhibits nitrogen fixation, affectingtotal nitrogen uptake in the legume host aswell as its contribution to the soil combinednitrogen content.

Abiotic stress activatescomplex signalling pathways

To cope with these stresses, plants haveevolved complex cell signalling pathwaysactivating metabolic functions anddevelopmental switches to permit theiradaptation to these conditions. The plantroot system, which is the primary site ofperception of environmental changes, isable to adapt its growth and architecture.This adaptation is a result of integratedevents occurring at all levels of organisation,from anatomical to morphological, andto cellular, biochemical, and molecular.Cellular responses to stress include changesin the cell cycle and cell division,modifications in the endomembrane systemand vacuolisation of cells, and changes incell wall architecture, all leading to enhancedstress tolerance of cells. At the biochemicallevel, plants alter their metabolism in variousways to accomodate environmental stresses,including the production of osmoregulatorycompounds.

The molecular events linking theperception of a stress signal with the genomicresponses leading to tolerance have beeninvestigated intensively in recent years,mainly in Arabidopsis. However, the universalnature of these mechanisms is still debated

because different strategies may be used bydifferent plants to cope with environmentalstresses. Insights into gene functions andregulatory control of biological processesthat are associated with stress responses inlegumes have been facilitated with thedevelopment of genomic resources andinformation for the two model legumespecies, Medicago truncatula and Lotusjaponicus (7, 8). For each of these tworeference legumes, extensive investmentsare being made to generate a full range ofgenomics resources, including sequencingthe entire genome and thus giving accessto a complete description of the responseat gene expression level. The increasingavailability of genetic and genomic data aswell as the high degree of synteny betweenlegume genomes has resulted in these twospecies becoming valuable models for themolecular genetic study of abiotic constraintshampering yield, since they are very closeto legume crops. Indeed, combininggenomic and biological knowledge aboutreference legumes that has a bearing onother food and feed legumes of majoreconomic importance, presents a majorscientific opportunity that can impact uponother less-easily handled crops.

Transcriptome analysis givespromising results

A critical step of the cell signallingpathways controlling stress responses involvestranscriptional regulation, generally mediatedby transcription factors that may governand coordinate the expression of large groupsof genes. In legumes, extensive sequencinghighlighted around 2,000 transcriptionfactors per genome, less than 1% of themgenetically characterised (9) and only six

Legumes provide more than one-thirdof mankind’s nutritional nitrogenrequirement and represent about25% of the world’s major crop production(6). Legumes can interact symbioticallywith nitrogen-fixing soil bacteria to fixatmospheric nitrogen, reducing productioncosts and environmental damage becausethey do not require soil fertilisers. Inaddition, the roles of legumes in the wholecrop rotation (for N cycle, soil structureand activities, disease and weed break, etc)benefit other subsequent non-nitrogenfixing crops. Therefore legumes play anessential role in sustainable agriculture.

Only 10% of arable land is consideredto have non-stressed soils, implying thatcrop growth on the remaining 90% of arableland is submitted to diverse environmentalstresses (3). Indeed, various abiotic stressesincluding salinity, drought, extremetemperatures (heat, freezing, chilling),waterlogging and mineral toxicities, reducethe yields of most major crops by morethan 50% (1). Abiotic stresses affect legumegrowth, yield and productivity as well asthe efficiency and function of the symbioticnitrogen fixation process. In general,exposure of plants to dehydration or osmoticstress causes a decline in photosyntheticactivity due to stomatal closure and reducedactivity of photosynthetic enzymes limitingcarbohydrate metabolism and plant growth.On a world scale, soil salinity is also aprimary cause of crop yield losses on morethan 20% of agricultural land and 50% ofcropland (4). High salt depositions in the

*Institut des Sciences Végétales – CNRS, Gif-sur-Yvette and Université Paris Diderot, France.([email protected], [email protected])



involved in abiotic stress tolerance (drought,salinity, extreme temperature). Thedevelopment of novel powerful tools fortranscriptome analysis increased considerablythe access to gene functions on a globalgenome-wide scale for the identificationof differentially expressed genes in responseto stress in legumes. These tools includesuppression subtractive hybridisation (SSH)library, super serial analysis of gene expression(SuperSAGE) for genome-wide quantitativegene expression profiling, array-basedtranscript profiling technologies (measuringexpression levels of tens of thousands of genesin parallel), and massive quantitative real timePCR (qRT-PCR) with specific in silicodesigned primers against known genes, suchas transcription factors (9). Recently, aMt16K microarray covering 16,086 tentativeconsensus sequences derived mainly fromapproximately 164,000 M. truncatula ESTscollected in the TIGR M. truncatula GeneIndex 5 (http://www.tigr.org/tdb/mtgi),was obtained. This Mt16K microarray wasused to monitor, for example, changes inthe transcriptome of dessication-sensitiveradicles of M. truncatula seeds at differentpoints in time, and therefore to investigateregulatory processes and protectivemechanisms leading to dessication tolerancein seeds (2). Another example is theapplication of SuperSAGE technology toprofile transcripts of drought- and salt-stressed roots and nodules from chickpeagiven access to a series of genes exclusivelyexpressed under both stresses, but not innon-stressed controls.

Novel regulatory genesrevealed by genomics

Thus, genomics and transcriptomicsprovide powerful tools to fully understandthe molecular basis of responses to abioticstresses in legumes as well as to the capacityof the plant to cope with environmentalstresses. This may lead consequently to theidentification of regulatory genes able tomodify crop behaviour under adverseconditions (as schematised in Figure 1). Asa recent example, expression analysis inmodel Medicago plants using a ‘dedicated’macroarray (containing selected genes fromSSH libraries), revealed novel regulatorygenes, including transcription factors,associated with reacquisition of root growthafter a salt stress (5). These selected genes

were classified into different functionalcategories depending on homologies toother species. Several stress-related genesinclude specific legume ESTs suggesting,therefore, the presence of novel pathwayslinked to abiotic stress responses in this family.Over-expression of one of the transcriptionfactor genes (MtZpt2-1) linked to recoveryprocesses in transgenic Medicago roots allowedthem to grow under salt stress conditionsand affected the expression of three putativetargets in a predicted manner: a cold-regulated A homolog, a flower-promotingfactor homolog and an auxin-inducedproline-rich protein gene. This establishesa role for this gene in the activation of aspecific genetic programme in theadaptation of legume roots to salt stress.

Combining genomics andgenotypic diversity

A large diversity of genotypes adapted toabiotic stresses was found in the glycophyteM. truncatula. This resource of diversity isused by breeders to manage abiotic stressesin their cultivation areas. Complex geneticsystems subject to interactions betweengenotype and environment controlled theplant traits for adaptation to environmentalstresses. Therefore, by using comparativegenomic approaches between thesegenotypes, the right complements of genesand alleles in a breeding programme could

be proposed. The understanding of thegenetic mechanisms involved in theseadaptive processes is particularly importantin cases such as drought or salt tolerancewhere few genes may drastically affect thebehaviour of related genotypes in a specificenvironment. Advances in genomic toolscombined with the exploitation of geneticdiversity from model legumes are allowingthe dissection of the genetic control ofcomplex traits. Determination ofphysiological tolerance mechanisms indiverse genotypes to various abiotic stressesin legumes may direct agronomicimprovement of legumes increasingproduction in a sustainable way. �

(1) Bray, E. A. et al. (2000). In: The biochemistryand molecular biology of plants, 1158–1249(Eds B. Buchanan et al.) The American Societyof Plant Physiologists, Rockville, USA.

(2) Buitink, J. et al. (2006). Plant J. 47, 735–750.

(3) Dita, M. A. et al. (2006). Euphytica 147, 1–24.

(4) Flowers, T. J. and Yeo, A. R. (1995). Aust. J.Plant Physiol. 22, 875–884.

(5) Merchan, F. et al. (2007). Plant J. 51, 1–17.

(6) O’Brian, M. R. and Vance, C. P. (2007).Plant Physiol. 144, 537.

(7) Pedrosa, A. N. et al. (2002). Genetics 161,1661–1672.

(8) Thoquet, P. et al. (2002). BMC Plant Biol. 2, 1.

(9) Udvardi, M. K. et al. (2007). Plant Physiol.144, 538–549.

Drought Salinity

Heat Freezing Chilling

Mineral toxicities

Waterlogging

ABIOTIC STRESSES

ADAPTATION

ARCHITECTUREGROWTH

SYMBIOTIC NODULES

REGULATORY GENESTRANSCRIPTION FACTORS

GENOMICTRANSCRIPTOMIC

APPROACHES

Mt16KMICROARRAYS

qRT-PCR

Figure 1. General view of the different steps linking genomics to the identification of key regulatory genesinvolved in abiotic stress responses in legumes. Exploiting genetic diversity in model legumes combined withtransgenic approaches will reveal regulatory mechanisms involved in legume adaptation to abiotic stress.


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Abiotic stresses, salinity, cold anddrought have the largest impact,causing yield losses of up to 50%in legumes. Evidence from various speciessuggests that abiotic stress tolerance is aquantitative, developmentally regulated,stage-specific phenomenon, and thattolerance at one stage of development maynot be correlated with tolerance at othersuch stages. This suggests that specific stagesshould be evaluated separately to developcultivars with stress-tolerance characteristics(1). Altogether, this implies that abiotic stresseshave an impact on a large number ofinteracting genes. Plant breeding thus facestwo questions: a) how to identify all the genes,and their allelic variations, that each confera better tolerance to stress and b) how tocombine all these genes in elite cultivars (3)?

A number of genes involved in abioticstress tolerance have been identified usingmolecular approaches (4, 6). However, itis very difficult to reveal their contributionto the plant phenotype in a givenenvironment, especially since these genesare often involved in more than one stress.Only the combination of molecular geneticapproaches such as Transmission genetics,Association genetics or Ecotilling allows abridge between molecular and genetic data.

The need for a model plantThe large number of gene x environment

x cultivated species combinations requiredfor legume breeding justifies the use oftaxonomically related model plants. Specialinterest in M. truncatula as a model plant forlegumes relies on its capacity to supportmolecular genetics programmes and itstaxonomic proximity with crops such as

alfalfa, pea, chickpea, lentil and fababean.Many molecular and genetic tools (cDNAand BAC banks, software, genetic maps)have been developed in conjuction withM. truncatula (5) and the sequence of itsexpressed genome is expected to be finishedin 2008.

One factor of major interest is thatM. truncatula grows spontaneously all aroundthe Mediterranean basin in very diverse,and sometimes hostile environments.Therefore, allelic variations responsible foradaptation to most abiotic stress could beexpected within M. truncatula naturalvariations. In addition to the existingcollections of M. truncatula (8, 2), a dedicatedcollection has been created based onTunisian natural populations growing indiverse eco-environmental conditions (7).

Molecular genetics ofM. truncatula salt tolerance

We first selected the genotypes Jemalongand F83005.5 which show contrastingphenotypes when submitted to salt stress(Figure 1). Using the M. truncatula geneticmap and the population of RecombinantInbred Lines (RILs) already developed, weidentified a number of Quantitative TraitLoci QTLs) involved in salt tolerance (Figure1). These QTLs underly genes whose allelicvariations are responsible for the observedphenotypic differences between these twogenotypes. But how are QTLs and the actualgenes connected? Transcriptomic approachesprovide important information about putativecandidate genes but only the cloning of the

alleles from one parent and its transfer tothe other will give the final proof. This map-based cloning of these QTLs is in progress.

Since the genes/alleles responsible forsalt-tolerant behaviour are not always thesame as in the plants analysed, this geneticapproach should be repeated using othercontrasting genotypes in order to identifymore QTLs/genes and evaluate their rolein the plant response to salt stress.

From models to cropsDue to the conservation of genome

organisation between related species, thelocation of a gene in a crop species couldbe deduced from model species studies andcould thus be used for marker- or genome-assisted selection of crops. However, itstill has to be established whether thefunction of a gene, and its allelic variations,are similar in different, even closely related,species. This research should be developedin the future. �

(1)Bayuelo-Jiménez, J. S. et al. (2002). CropScience 42, 2184–2192.(2) Ellwood, S. R. et al. (2006). Theor. Applied.Genet. 112, 977–983.(3) Flowers, T. J. et al. (1999). Journal ofExperimental Botany 51, 99–106.(4) Mahajan, S. and Tuteja, N. (2005). Arch.Biochem. Biophys. 444(2),139–158.(5) Medicago truncatula handbook:http://www.noble.org/MedicagoHandbook/(6) Merchan, F. et al. (2007). The Plant Journal51, 1–17.(7) Lazrek, F., Huguet, T. and Aouani, M. E. Inpreparation.8) Ronfort, J. et al. (2006). BMC Plant Biology6, 28.

*Laboratoire Symbioses et Pathologies desPlantes (SP2), INP-ENSAT, Castanet Tolosan,France. ([email protected])

Integrating molecular and genetic data in legumesIntégration des approaches moléculaires et génétiques chez leslégumineusesby Thierry HUGUET*

Figure 1. Contrasting tolerance to NaCl (45mM) of M. truncatula Jemalong (left) and F83005.5 (right);M.truncatula genetic map based on LR5 population of RILs. Vertical lines indicate QTL locations involved in thereduction of stem growth (1), reduction in the number of leaves (2), reduction of root growth (3).

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Benefits of inoculation under abioticstress in the Mediterranean regionL’intérêt de l’inoculation sous stress abiotique en région méditerranéenneby Carmen VARGAS* and Mohamed Elarbi AOUANI**

*University of Seville, Spain. ([email protected])

**Centre de Biotechnologie, Technopole deBorj Cedria, Tunisia and NEPAD/North Africa Biosciences Network (NABNet), National Research Centre, Cairo, Egypt.([email protected])

If appropriately managed, and especiallyunder abiotic stress (drought, salinity),microbial biofertilisers can improvefood legume yield and soil fertility andreduce pollution by inorganic fertilisers. Ingeneral, three types of inoculants can helplegumes overcome stressful environments:

• Efficient and stress-tolerant symbioticrhizobial strains, used when native populationsare scarce and/or inefficient. The Aquarhizproject1 targeted four south Mediterraneancountries (Algeria, Egypt, Morocco,Tunisia) to study the inoculation of fababean, chickpea and common bean underdrought and/or saline conditions. Severaltrials showed a positive effect of inoculation,with variation according to countries andcrops but also according to strains. At someexperimental sites the inoculation had noeffect or the increase in yield was notsignificant. In the case of faba beans inMorocco, soils are naturally hosting efficientlocal strains. However, in several other cases,sowing seeds with addition of these stress-tolerant rhizobium strains enhanced nitrogenfixation activity and improved yield of grainlegumes. The symbiotic systems which areinhibited by abiotic stress are well known:oxygen permeability, shortage of watersupply to nodules from the phloem,overproduction of reactive-oxygen species,feedback inhibition involving shoot Nstatus. However the mechanisms underlyingthe stress tolerance of symbiosis remainunclear. Candidate systems from the plant

(aquoporins, carbonic anhydrase) or thebacterial (osmoprotectants, bacteroidalrespiration) side should be investigated inorder to select successful combinations oflegume and microsymbionts better adaptedto stress.

• Arbuscular mycorrhizal fungi (AMF)colonise root systems and form symbiosescalled mycorrhiza with almost all legumes.By exploring a greater volume of soilthrough the fungal hyphae, mycorrhizaenhance nutrient (particularly phosphorus)and water uptake from the soil, therebyenhancing plant growth under drought stress.

• Plant Growth Promoting Rhizobacteria(PGPR), include a diverse group of free-living soil bacteria that can stimulate plantgrowth by a number of direct or indirectmechanisms. Interestingly, many strainsisolated in the Aquarhiz project are free-living non-rhizobial species with promisingapplications as inoculants.

Inoculation is limited anddecreasing

Although inoculation is a well developedagricultural practice in Australia, Canada,USA and Latin America, it is not commonin Europe and Africa. The status of legumeinoculation in the Mediterranean region andthe related research strategies were discussedat a FABAMED2 workshop organised atRabat in February 2005, following the firstGeneral Meeting of the Aquarhiz project.It was shown that legume inoculation withrhizobia was limited and decreasing in theregion, whereas the use of N fertilisers wasincreasing, especially at Egypt.

The workshop participants concludedthat the symbiosis should be taken intoaccount in research activities and in the

technical messages disseminated to farmersto improve legume production in theMediterranean region, but some key pointsshould be considered:

- The inoculation is not the uniquesolution as shown in past experience, anda systematic diagnosis is required torationalise where and when inoculantapplication can have an impact.

- Active rhisophere and an efficientrelationship between enhanced cultivarsand the best adapted rhizobia, either presentin the soil or added as inoculant, is a keycomponent to boost N fixation and rootgrowth leading to better tolerance of abioticstress.

- Inoculation should be associated withappropriate crop and soil management.Plant genotype and nutrient conditions arekey factors influencing inoculation success.

- Progress in inoculation technology isrequired, i.e. second generation inoculants(rhizobial plus mycorrhiza or PGPR).

- Inoculant production is an easy and cheaptechnology that could be scaled-up withinstitutional support.

- Farmers’ adoption of the technology willrequire participatory field research andappropriate technological packages. Theseactivities have been initiated within theframework of the Aquarhiz project. �

1FP6 project www.grainlegumes.com/aquarhizand Grain Legumes 39, 23.2FABAMED is a network of scientists interestedin N fixation in the Mediterranean Region. Setup in 1994, it meets on a regular basis forinformation exchange, workshops and thepreparation of joint projects.

Faba beans with inoculation (left of photo) andwithout inoculation (right of photo), grown in sandysoils in Beheira in Egypt.

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ABIOTIC STRESS

*ICARDA, Aleppo, Syria. ([email protected])

Abiotic stress breeding at ICARDAAmélioration variétale et stress abiotique à l’ICARDAby Michael BAUM*

The mission of the InternationalCenter for Agricultural Researchin the Dry Areas (ICARDA) is tomeet the challenges of dry-areaenvironments, which are harsh, stressfuland variable. ICARDA has both global andregional mandates. It has a globalresponsibility for the improvement of threeimportant food crops – barley, lentil, andfaba bean. ICARDA's regional responsibilitywithin Countries of West Asia and NorthAfrica (CWANA) focuses on theimprovement of wheat, chickpea, forageand pasture crops.

ICARDA holds the largest gene bankin the Mediterranean region, with about133,000 accessions, which representsapproximately 20% of the germplasm inCGIAR centres. Particularly important arethe landraces and wild relatives that haveevolved under harsh conditions overmillennia. About 70% of ICARDA’scollections are now geo-referenced. Thiscollection-site information combined withclimatic layers in Geographical InformationSystem (GIS) allows targeted collection anda rapid exploitation of accessions withtolerance to drought and heat to meet theanticipated effects of climate change.ICARDA has been sharing these resourcesfreely with partners all over the world.On average, the Center distributes 35,000samples per year. Overall, the shift fromcollection and ex situ conservation of plantgermplasm to its characterisation, evaluationand documentation will help to utilise thebiodiversity held at ICARDA.

Using modern technolgiesICARDA collaborates with advanced

research institutes (ARIs) and the NationalAgricultural Research Systems (NARS) in

using modern technologies for its cropimprovement programmes. Emphasis isgiven to the identification and exploitationof genetic resources for improved stressresistance and water-use efficiency. DNAmolecular marker techniques allowconstruction of linkage maps for crops.Together with statistical techniques theselinkage maps can be used to locate andestimate phenotypic effects of quantitativetrait loci (QTL) and the genes responsiblefor the expression of agronomic traits. Fora homozygous population derived from across with parents contrasting in responseto, for example, water, QTL analysis revealsthe approximate map location of lociassociated with performance under drylandconditions. This is then amenable tomarker-assisted selection using DNAmarkers flanking the identified QTLs.

Lentil and chickpeaimprovement

Radiation–frost injury is an importantabiotic constraint to lentil production inWest Asia. If cold tolerance of lentil couldbe improved, its yield in low rainfall areascould be increased by early planting.Recombinant inbred lines of a lentil crosswere tested in northern Syria for plant frostinjury levels and genotyped with 254 DNAmarkers. DNA markers were identifiedlinked to the locus for radiation–frosttolerance trait and Fusarium wilt resistance.These are proving useful in lentilimprovement. Instead of selecting forradiation–frost tolerance and Fusarium wilt,breeders can select for the resistance markeralleles.

Chickpea is an important dryland cropsown in spring in WANA. Its yield andwater-use efficiency can be nearly doubledby advancing its date of sowing to winter.Ascochyta blight, a fungal disease attackingchickpea, has developed into a major threat

for winter-grown chickpea. DNA markershave been used to characterise the availablepathotypes of the fungus. This allows thedevelopment of geographical distributionmaps of the pathogen and the deploymentof effective host-plant resistance genes. Fortagging host-plant resistance genes inchickpea, microsatellite based markers havebeen developed in collaboration with theUniversity of Frankfurt. Host-plantresistance for Ascochyta blight is beingmapped in several populations and geneticbackgrounds.

BiosafetyICARDA is also exploring the possibility

of using genetic transformation to achieveimproved tolerance to drought and otherabiotic stress resistance. ICARDA alsoactively promotes the development andestablishment of national and regionalbiosafety regulations. So far, Egypt andSyria are the countries in the region thathave established biosafety regulations. Inseveral other countries the preparations forthe development of biosafety regulationsare underway. ICARDA’s policy is guidedby international biosafety practices and theregulations of the Syrian Arab Republicfor the development and deployment ofGMOs. �

Field of chickpeas at Ain Temouchet, Algeria inApril 2007.

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Where we go from here – concluding remarks of theabiotic stress in legumes workshop Quelles perspectives pour la suite – conclusion du séminaire sur lestress abiotique chez les légumineusesby Martin CRESPI*, Anne SCHNEIDER**, Thierry HUGUET*** and Mohamed Elarbi AOUANI****

Maintaining or improving cropproductivity under conditions ofabiotic constraint in the field isone major concern in many areas in theworld where legumes are grown. Abioticstresses refer to different environmentalfactors: water deficit, high temperatures,saline stress, mineral deficiency, frost,chilling, and others, which can occur atdifferent levels and development stages. Asshown in the Tunis workshop1, it is veryuseful to integrate different disciplinaryexpertise to apprehend the complexity oftolerance and susceptibility traits.

From whole plant to genomicapproaches, a strategy toexploit genetic diversity

The whole plant approach allows keyvariables of the plant/genotype interactionthat maintains yield under stress to be assessed.Environmental signals and plant physiologicalmechanisms activated at different growthstages can be useful to determine theirhierarchy and relevance in the physiologicalprocesses involved. Adapted agronomicmanagement techniques are needed to copewith abiotic stresses in the field. However,modelling the eco-physiological interactionsbetween plants and the environment,considering the particular aspects of legumephysiology, will enhance the basicunderstanding of mechanisms for furtheragronomic improvement.

In legumes, the symbiotic bacterial partner,the rhizobium, and other components ofthe rhizosphere have key roles in the

agronomic performance of these crops, andshould be taken into account in the analysisof plant response. Improving symbiosisunder stress conditions will contribute tothe development and use of legumes aspioneering crops since dry or saline soilsare also poor in combined nitrogen.

Parallel development of genomicapproaches will reveal molecularmechanisms involved in the regulation ofthe plant physiological responses. Exploitingthe data obtained for different plant species(particularly different legumes) willhighlight key molecules and genes involvedin stress responses. These approaches shouldevolve from the initial description levelunder specific conditions to more elaboratescreenings defining traits and genes involvedin crop adaptation strategies. Targeted cropstrategies should focus on the enhancementof tolerance to abiotic stress during grainfilling or on rapid soil coverage in specificsystems by controlling germinationpotential and initial root growth.

In addition, exploiting genetic diversityis crucial to obtain both contrasted typesand sources of tolerances. Genetic resourcesare in fact mutants already selected byspecific environmental conditions, possiblymaking them more suitable for fieldconditions than laboratory mutants.

Interdisciplinary approachesshould be developed

Integrating molecular and genetic datainto eco-physiological models is requiredin order to define the regulatorymechanisms involved in the control of plantgrowth and development under abioticstress constraints and to create novel elitecultivars. This will bring new perspectives

to the development of novel cropmanagement techniques and to theidentification of crucial breeding targets.

The general approaches favoured in orderto make progress in this area were discussedamong Tunis workshop participants. Theneed for interdisciplinary approaches toenhance knowledge in abiotic stress responsesand tolerance was agreed upon. Plantphysiology expertise should be reinforcedparticularly to establish common referencesand protocols in case studies: how to applystress, what sort of stress and its level ofintensity, what type of plant response istargeted (e.g. survival or adaptation). Geneticmodels will provide information on plantresponses at different levels that can be appliedin crop species although the specificity ofthese responses according to species or evenlines should also be considered. Two differentresearch strategies were debated: one focussingon the search for few key genes whosemodulation would provide a breakthrougheffect on crops or a more integrative approachaiming to dissect the complex mechanismsinvolved in stress responses. Both willcontribute to the development of improvedvarieties adapted to cropping systems.

In spite of strong debate, specific pointswere unanimously agreed: the kindhospitality of the Tunisian hosts, the interestin participating in interdisciplinaryworkshops and the need to structurecollaborations between experts fromdifferent fields to increase the relevance ofthe strategies in crop management.

1‘Abiotic stresses ain legumes’ (Tunis, 22–24March 2007), a scientific workshop organised atthe initiative of the Grain Legumes IntegratedProject (GLIP) which is supported by theEuropean Commission from 2004 to 2008.

*CNRS-ISV, Gif sur Yvette, France**AEP, Paris, France***INRA-ENSAT, Toulouse, France****CBBC-LILM, Hammam Lif, Tunisia


Dynamics and prospects for grain legumes in theEuropean feed marketDynamiques et prospectives pour les protéagineux sur le marchéeuropéen de l’alimentation animaleby Anne SCHNEIDER*, Frédéric PRESSENDA** and Katell CRÉPON***

analysed, only half the apparentconsumption was used for compound feed(194,000 t vs. 400,000 t) and according tothe model, using the observed market price,the potential use could reach 730,000 t(Figure 1). The maximum potential usecould reach 1.05 Mt if the market pricewas halved. Interviews with Germancompound feed manufacturers highlighteddiverse reasons for the current under-useof peas: (i) negative attitude of their clientstowards pea, (ii) interest in pea prices butproblems of year-round availability for somemanufacturers, (iii) higher pea prices inGermany because of pea exports to theNetherlands caused by an attractive market.

Analysis of the shadow prices2 of peashows that nearly all formulas (except thatof ruminants) were able to incorporate peaduring the period July 03 to June 04, themarket price of pea being lower than theshadow price. Pea is particularly interestingfor piglet, fattening pig and sow diets, butalso for broiler and laying hen diets.

the EU highly dependent on imports. Thedeficit in Materials Rich in Protein (proteincontent >15% of dry matter) in the EU27amounts to 73% and two-thirds of this isoffset by imports of soyabean seed and mealwhich amount to 35 Mt annually (2).

In recent GLIP1 analyses, it has beenshown that with current seed nutritionalcomposition, technical constraints and marketprices, more European grain legumes couldbe utilised for feed than the volumes currentlyutilised (Pressenda, personal communication,March 2007). When the production levelsof pea and faba bean and their apparentutilisation (compound feed production andon-farm feed uses) are analysed, two typesof countries can be defined:

(i) producing countries: France,Germany, UK, Denmark, Czech Republic;

(ii) importing countries: Spain,Netherlands and Belgium.

In Germany, when the potential use ofpeas and faba beans in compound feeddestined for different animal species was

*AEP, Paris, France. ([email protected])

**CEREOPA, Paris, France.([email protected])

***UNIP, Paris, France. ([email protected])

Until now, the animal feed industryhas been the predominant outletfor European grain legumes evenif there are recent expanding added valuemarkets: food exports and ingredients forfood, feed or non-food uses.

About 85% of European pea productionand 58% of faba bean production was usedby the feed industry in 2003/2004, especiallyin France, Germany and Spain. In the UK,the percentage was smaller (75% feed and25% food uses). At least 3.5 million tonnes(Mt) of grain legumes are used by the animalfeed industry (1), and there are also nichemarkets such as pigeon feed or pet food(stable demand) as well as on-farm feed uses.

Legume grains are energy- and protein-rich raw materials used for different animalswith different nutritional requirements:monogastrics and ruminants constitute themajor segment, fish diets a minor one. Fababean which is richer in protein than pea isinteresting for poultry. Lupins are moresuitable for ruminants than pea and fababeans. In France in the last 12 years peashave been used mainly for pig compoundfeed (90%), and in smaller amounts forpoultry (8%) (1).

Current use could be muchlarger

Less than 3% of the 130 Mt of the EUcompound feed is from protein crops. Thedemand for grain legumes from thecompound feed industry greatly exceedsthe level of home production, which makes

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