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  • Omics and modelling approaches for understanding regulation of asymmetric cell divisions in arabidopsis and other angiosperm plants

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  • Omics and modelling approaches for understanding regulation of asymmetric cell divisions in arabidopsis and other angiosperm plants
    View larger version In this window In a new window Download as PowerPoint Slide Fig 4 Schematic representation of the primary root of arabidopsis indicating the developmental progression along the primary root axis pre initiation events leading to asymmetric cell division ACD i Priming stage in basal meristem region in which generation of an auxin maximum yellow in the protoxylem cells alongside oscillating gene expression in the region leads to formation of prebranch sites in the adjacent xylem pole pericycle cells XPPs ii These XPP cells have the competence to form lateral roots and pass through a developmental window to become specified founder cells FCs iii Migration of nuclei towards the common cell wall indicates FCs preparing to undergo asymmetric cell division iv Anticlinal asymmetric cell division of FCs occurring in the differential zone of the root giving rise to two small daughter cells deep purple and two larger flanking cells pale purple also representing the stage I primordium v Following anticlinal division the axis of division shifts by 90 with the central core of small daughter cells represented by deep purple in iv dividing in an outward manner giving rise to equally sized daughter cells with different identities orange and green also representing stage II primordium A selection of key players regulating these processes is indicated for the distinct steps View larger version In this window In a new window Download as PowerPoint Slide Fig 5 Procambium and cambium asymmetric divisions A Diagram of a cross section of primary and secondary stems Primary stems have vascular bundles where asymmetric divisions of procambium also called fascicular cambium produce cells for procambium phloem and xylem When the stem transitions from primary to secondary growth interfascicular cambium is induced which allows cambium phloem and xylem to form continuous rings around the stem B Schematic showing the known interactions in the PXY TDR signalling pathway across the cell types in the stem View larger version In this window In a new window Download as PowerPoint Slide Fig 6 General schemes for asymmetric divisions ACD in stomatal development A Stomatal development in the grasses Cells are arranged in linear files along the length of the leaves In alternate files at the base of the leaf asymmetric divisions generate guard mother cells GMCs blue GMCs signal to the neighbouring cell files inducing the migration of subsidiary mother cell SMC nuclei towards the point of contact with the GMCs and the subsequent asymmetric and oriented cell division of the SMCs to form the subsidiary cells This is facilitated by the polarized proteins PAN1 and PAN2 yellow lines B In dicots such as arabidopsis stomatal precursors are not arranged in files and an additional step precedes the formation of GMCs The initiation of the stomatal lineage is an asymmetric division of a meristemoid mother cell MMC white to produce a meristemoid green and a stomatal lineage ground cell SLGC A meristemoid can divide again in an amplifying division renewing itself and producing another SLGC lower pathway and

    Original URL path: https://aob.oxfordjournals.org/content/113/7/1083.figures-only?sid=b5a6bc25-5f3c-4c5f-a5e2-76b73d864083 (2016-02-18)
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  • Lathyrus diversity: available resources with relevance to crop improvement – L. sativus and L. cicera as case studies
    L sativus and L cicera Conclusions Efforts for improvement of L sativus and L cicera should concentrate on the development of publicly available joint core collections and on high resolution genotyping This will be critical for permitting decentralized phenotyping Such a co ordinated international effort should result in more efficient and faster breeding approaches which are particularly needed for these neglected underutilized Lathyrus species Key words Diversity genetic resources Lathyrus sativus L cicera Fabaceae legumes plant breeding protein crops The Author 2014 Published by Oxford University Press on behalf of the Annals of Botany Company All rights reserved For Permissions please email journals permissions oup com Related articles ContentSnapshots Content Snapshots Ann Bot 2014 113 6 i iii doi 10 1093 aob mcu079 Extract Full Text HTML Full Text PDF Previous Next Article Table of Contents This Article Ann Bot 2014 113 6 895 908 doi 10 1093 aob mcu024 First published online March 12 2014 Abstract Free Free Figures Free Full Text HTML Free Full Text PDF Free All Versions of this Article mcu024v1 113 6 895 most recent Classifications Invited Review Services Article metrics Alert me when cited Alert me if corrected Alert me if commented Find similar articles Similar articles in Web of Science Similar articles in PubMed Add to my archive Download citation Request Permissions Responses Submit a response No responses published Citing Articles Load citing article information Citing articles via CrossRef Citing articles via Scopus Citing articles via Web of Science Citing articles via Google Scholar Google Scholar Articles by Vaz Patto M C Articles by Rubiales D Search for related content PubMed PubMed citation Articles by Vaz Patto M C Articles by Rubiales D Agricola Articles by Vaz Patto M C Articles by Rubiales D Related Content Related articles in this journal Load related web page information Share Email this article CiteULike Delicious Facebook Google Mendeley Twitter What s this Search this journal Advanced Current Issue February 2016 117 2 Alert me to new issues The Journal About this journal Annals of Botany Collections AoB article attracts media coverage We are mobile find out more Journals Career Network Published on behalf of The Annals of Botany Company Impact factor 3 654 5 Yr impact factor 4 338 Eigenfactor 0 02603 Rank 10 200 SCImago Score 1 461 Rank 124 1873 Chief Editor Professor J S Pat Heslop Harrison View full editorial board International Review Board For Authors Submitting a manuscript online Self archiving policy Instructions for authors Low Rate Open Access Fees Open access options for authors visit Oxford Open Visit HighWire Press 3hWaciBYRk30rSOQ7UOpP6viAxsZnEle true Looking for your next opportunity Looking for jobs Alerting Services Email table of contents Email Advance Access CiteTrack XML RSS feed Rights Permissions This journal is a member of the Committee on Publication Ethics COPE Corporate Services Advertising sales Reprints Supplements Widget Get a Widget Most Most Read Calcium in Plants Pollen Tube Distribution in the Kiwifruit Actinidia deliciosaA Chev C F Liang Pistil in Relation to its Reproductive

    Original URL path: https://aob.oxfordjournals.org/content/113/6/895.abstract?sid=b5a6bc25-5f3c-4c5f-a5e2-76b73d864083 (2016-02-18)
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  • Lathyrus diversity: available resources with relevance to crop improvement – L. sativus and L. cicera as case studies
    2000 based on quality analysis and by Hanbury et al 1999 based on agronomic testing Those lines of Mediterranean European origin were consistently higher yielding with much larger seeds and later phenology Hanbury et al 1999 They also had lower ODAP content Abd El Moneim et al 2001 Preference for larger seeds in this area is common to other grain legumes such as lentil Lens culinaris chickpea Cicer arietinum and faba bean Vicia faba and is a product of human selection Chowdhury and Slinkard 2000 Similar studies on field evaluation of grass pea landraces but in a more restricted germplasm study where high variability in morphological and agronomical traits was detected were also performed with Chilean Tay et al 2000 Ethiopian Tadesse and Bekele 2003 a b Italian Tavoletti et al 2005 Indian Kumari 2001 Spanish De la Rosa and Martín 2001 and Slovak germplasm Benková and Záková 2001 In the majority of these studies diversity among and within populations has been detected for several of the characterized traits Table 3 indicative of high breeding potential in these materials View this table In this window In a new window Table 3 Examples of Lathyrus diversity studies in morphological traits Diversity analysis through biochemical and molecular markers Biochemical and molecular markers can be used to better document the organization of genetic diversity between possible parental materials of new breeding programmes The high agronomical and morphological diversity within L sativus germplasm is also found at the biochemical and molecular level Considerable genetic diversity as revealed by isozymes and molecular markers exists in L sativus throughout the world Table 4 These markers are normally very efficient in distinguishing among different L sativus genotypes However it was not always possible to associate genetic diversity with morphological or geographical diversity Yunus et al 1991 Tadesse and Bekele 2001 Belaid et al 2006 Vaz Patto et al 2011 The lack of correlation between genetic diversity and the region of origin supports the idea that the natural distribution of L sativus has been completely obscured by cultivation View this table In this window In a new window Table 4 Examples of Lathyrus diversity studies in biochemical and molecular markers Chowdhury and Slinkard 2000 using a wide L sativus germplasm collection managed however to associate different levels of genetic diversity measured by isozymes with the different geographical origins These authors identified the Near East and North Africa regions as those with the most isozyme variability suggesting that the centre of diversity for L sativus was this general area Also using a worldwide collection of L sativus accessions from several different geographical origins and the seed proteins albumins Przybylska et al 1998 and globulins Przybylska et al 2000 it was possible to separate two groups of L sativus accessions white seeded with large seeds originating mainly from Europe and North Africa and coloured seeded with relatively small seeds originating mainly from Asia and Ethiopia Nevertheless in a restricted study using only Southern Italian L sativus germplasm seed storage proteins were revealed to be unsuitable for detecting any variability among the studied landraces Lioi et al 2011 This may be an indication of the high level of genetic affinity among these landraces collected from a restricted geographical region PCR based molecular markers such as randomly amplified polymorphic DNA RAPD and intersimple sequence repeat ISSR markers have also proven to be efficient in distinguishing between different L sativus accessions and in assessing the within species genetic variability Chtourou Ghorbel et al 2001 Belaid et al 2006 Barik et al 2007 The presence of considerable intrapopulation variation among the L sativus accessions revealed in many of these diversity studies Chowdhury and Slinkard 1997 Gutierrez Marcos et al 2006 was greater than would have been expected given the predominantly autogamous breeding system of L sativus In fact although L sativus appears to be autogamous outcrossing rates as high as 36 have been recorded Rahman et al 1995 Chowdhury and Slinkard 1997 Gutierrez Marcos et al 2006 which have implications in breeding and germplam maintenance Tavoletti and Iommarini 2007 evaluated the genetic diversity of a collection of L sativus populations collected in central Italy using AFLPs Two main clusters were found one included large seeded populations from farms and the second included small seeded populations cultivated in market oriented farms AFLP markers have also been used more recently in a Southern Italian collection of L sativus and even though the detected polymorphism was low these populations were completely discriminated using 12 AFLP primer combinations Lioi et al 2011 The genetic diversity of a collection of Iberian L sativus germplasm was also studied using AFLPs as a first step towards the selection of appropriate parental lines for the establishment of a disease resistant cross breeding scheme Vaz Patto et al 2011 Molecular markers can also be developed using publicly available DNA sequencing data Expressed sequence tags ESTs in public databases and cross species transferable markers are considered to be a cost effective means for developing sequence based markers for less studied species Ellwood et al 2008 Both approaches have been applied to Lathyrus sp with variable achievements Molecular markers developed for closely related legume species have been shown to be transferable to L sativus and L cicera Almeida et al 2014 These included genomic and expressed sequence tag microsatellite gSSR and EST SSR and intron targeted amplified polymorphic ITAP markers and were successfully used to discriminate within L cicera and L sativus accessions Shiferaw et al 2012 using information on EST SSRs derived from Medicago truncatula and also on publicly available NCBI database L sativus EST sequences developed and validated polymorphic markers that were used successfully for exploring the genetic diversity of Ethiopian grass pea accessions Lioi et al 2011 developed SSR markers from publicly available EMBL database L sativus and Lotus japonicus cDNA sequences and used them to study Southern Italian L sativus accessions In this case accessions were grouped into two clearly distinguishable clusters following a geographical pattern but not consistent with the morphological data AFLP or SSR based clustering If we take into account the presence of polymorphism in the studies where this comparison could be performed it can be concluded that more informative markers for genetic diversity studies were developed directly from L sativus sequences than were transferable from M truncatula or Lotus japonicus More recently polymorphic EST SSRs were developed from L sativus sequence information available on a public database NCBI database Sun et al 2012 and from an enriched grass pea genomic library Lioi and Galasso 2013 as additional resources for grass pea genetic studies but they are not yet exploited in diversity analysis Diversity on quality traits Several preliminary studies to establish quality breeding approaches in Lathyrus sp resulted in characterization of the quality diversity of germplasm collections e g Granati et al 2003 Lathyrus species are protein rich legumes the development of which into important food legumes has been hindered by the presence of ODAP which if consumed in large quantities for extended periods can cause irreversible paralysis Lambein and Kuo 2009 The reduction in ODAP levels in L sativus breeding has been the emphasis for a long time Kumar et al 2011 Girma and Korbu 2012 Hillocks and Maruthi 2012 No L sativus or L cicera accession is ODAP free although in several lines the ODAP content can be significantly low This appears to be species related since the average ODAP content of L cicera is generally lower than that of L sativus Hanbury et al 2000 Abd El Moneim et al 2001 Kumar et al 2011 Variation of ODAP content in a range from 0 02 to 2 59 has been reported in L sativus Granati et al 2003 Tadesse and Bekele 2003 a Grela et al 2010 2012 Piergiovanni et al 2011 and from 0 09 to 0 49 in L cicera seeds Granati et al 2003 Sánchez Vioque et al 2009 Selection for high yield and low ODAP can be practised simultaneously for L sativus improvement Most of the initial progress in the development of cultivars low in ODAP was by direct selection from landraces and lines with a worldwide origin and several improved grass pea cultivars have been released as the result of various national and international breeding initiatives Ali Bar Ceora Gurbuz 1 Wasie Prateek Mahateora Ratan Bari Khesari 1 and 2 and Bina Khesari 1 all with an ODAP content 0 1 summarized by Abd El Moneim et al 2001 Kumar et al 2011 Similarly improved cultivars with low ODAP have been released such as Chalus Hanbury and Siddique 2000 This strategy of prioritizing reduction of ODAP content in breeding programmes is under debate today First although a number of cultivars with low ODAP have been released the long term results of these efforts are frequently questioned because ODAP content is highly influenced by climatic and edaphic conditions with strong genotype environment effects Fikre et al 2011 Jiao et al 2011 Girma and Korbu 2012 Water stress can double the toxin content in the plant Hanbury et al 1999 and there are indications that zinc fertilization can reduce the toxin accumulation Lambein et al 1994 although the mechanism by which the ODAP content may be reduced by added zinc is not known Abd El Moneim et al 2010 This long term breeding priority did not take into consideration that ODAP in itself does not seem to be a problem when grass pea is consumed as part of a balanced diet in which case grass pea is harmless to both humans and animals Lambein and Kuo 2009 Also risks of overconsumption can be reduced by the fortification of grass pea with cereals rich in sulfur amino acids and condiments rich in antioxidants such as onion garlic and ginger Getahun et al 2003 2005 In addition to this seeds can be partly detoxified by various food processing methods as reviewed by Kumar et al 2011 Therefore it seems clear that the widespread school of thought held 50 years ago of the vital need to reduce the ODAP content in Lathyrus seeds by breeding does not exist today Even with the possibility of its toxicity we should not neglect the potential benefits of ODAP For instance there is the prospect of using ODAP as a haemostatic agent during surgery Lan et al 2013 ODAP is not only produced by several Lathyrus sp seeds it is also present in the longevity promoting ginseng root Kuo et al 2003 where under the name Dencichine it is known for its haemostatic property to stop bleeding Lan et al 2013 In addition there is the hypothesis that nitriles are the causative agents of neurolathyrism rather than ODAP Llorens et al 2011 However nitriles too even though they are toxic can have some benefits For instance β aminopropionitrile β APN inhibits the cross linking of collagen and is the cause of osteo lathyrism but has a number of pharmacological applications β APN has the potential for the control of silicotic pulmonary fibrosis Levene et al 1967 for the control of unwanted scar tissue in humans Harrison et al 2006 and for diminishing the metastatic colonization potential of circulating breast cancer cells Bondareva et al 2009 β APN is a reagent used as an intermediate in the manufacture of β alanine and pantothenic acid Most reports on β APN refer to L odoratus Genetic variation for content in other Lathyrus germplasm has not been explored Another recent paradigm shift in the perception of the L sativus research is its content of homoarginine which is an alternative substrate for nitric oxide biosynthesis Rao 2011 Nitric oxide is well recognized for its role in cardiovascular physiology and general well being and thus a daily dietary intake of homoarginine through small quantities of L sativus may have advantages and deserves to be exploited The activation of protein kinase C PKC by ODAP adds a new dimension for investigating its therapeutic potentials in such areas as Alzheimer s disease hypoxia and the long term potentiation of neurons essential for memory Rao 2011 Genetic variation for homoarginine content in L sativus germplasm has been identified Piergiovanni and Damascelli 2011 Also Lathyus spp have potential for use as functional foods as the antioxidant activity of their polyphenols is higher than that of other legumes such as chickpea lupin Lupinus sp and soybean Glycine max Pastor Cavada et al 2009 b Another potential beneficial application of L sativus seeds it to ameliorate diabetic symptoms as they possess glycosylphosphatidylinositol with insulin mimetic activity Pañeda et al 2001 Diversity in stress resistance Many more reports exist on the biotic stress resistance evaluation of Lathyrus germplasm collections than on abiotic stress evaluations Lathyrus sativus and L cicera accessions of Iberian origin have been screened for resistance against powdery mildew and rust fungi Vaz Patto et al 2006 a b 2007 2009 Vaz Patto and Rubiales 2009 2014 and against the parasitic weed Orobanche crenata Fernández Aparicio et al 2009 2012 Fernández Aparicio and Rubiales 2010 identifying a wide range of levels of resistance Moderate levels of resistance to powdery mildew in L sativus have also been reported in India and Syria Campbell et al 1994 Robertson and Abd El Moneim 1996 Asthana and Dixit 1998 Powdery mildew is among the major diseases affecting L sativus Campbell et al 1994 and L odoratus crops Cook and Fox 1992 and rusts are important diseases of L sativus in north western Ethiopia Campbell 1997 However insufficient information is often available on the identity of the fungus Powdery mildew that infects Lathyrus is believed to be mainly Erysiphe pisi but it might be that several other species are able to infect Lathyrus sp as recently found in pea Fondevilla et al 2013 The existence of specialized forms and races is still unclear but a different ability to infect different plant species has been reported Cook and Fox 1992 reported that a strain of E pisi collected on L odoratus was able to infect faba bean but not pea whereas a different strain collected on L latifolius was able to infect pea and faba bean Similarly rust in Lathyrus sp is believed to be caused by both Uromyces pisi and U viciae fabae Barilli et al 2011 2012 Resistance to or escape from the parasitic weed O crenata has also been identified in L sativus and L cicera germplasm Fernández Aparicio et al 2009 2012 Fernández Aparicio and Rubiales 2010 High levels of resistance to O crenata have been reported in the species L ochrus and L clymenum Sillero et al 2005 Other relevant reports include resistance to Mycosphaerella pinodes Robertson and Abd El Moneim 1996 Gurung et al 2002 Fusarium oxysporum Benková and Záková 2001 and Cercospora pisi sativae Mishra et al 1986 in L sativus germplasm to Pseudomonas syringae pv syringae in L cicera germplasm Martín Sanz et al 2012 and to the northern root knot nematode Meloidogyne hapla in L latifolius L sylvestris and L hirsutus Rumbaugh and Griffin 1992 All these reports are summarized in Table 5 View this table In this window In a new window Table 5 Examples of Lathyrus diversity studies in biotic stress resistance In relation to Lathyrus abiotic stress resistance screening the lack of methodologies to identify resistant genotypes has hampered the proper exploitation in breeding of Lathyrus sp As a result knowledge of the mechanisms underlying this resistance to environmental injuries is also missing The effects of drought and salt stress on different Lathyrus sp morphological and physiological traits have been studied with the objective of developing the missing efficient discrimination methods applicable to large germplasm screenings Using a critical salt induced stress treatment or the chlorophyll a fluorescence transient several L sativus salt and drought resistant genotypes respectively have been identified Talukdar 2011 Silvestre et al 2014 Previous Section Next Section PROSPECTS FOR LATHYRUS DIVERSITY ANALYSIS AND USE IN BREEDING Conserved plant genetic resources are essential to meet the current and future needs of crop improvement programmes However progress in Lathyrus breeding has been slow due to the dispersal of the few available resources and evaluation efforts among several scattered germplasm collections plus the modest molecular and biotechnological breeding tools currently in existence More efficient and faster breeding approaches are needed on this neglected but promising underutilized species Marker development for diversity analysis and for marker assisted selection Although there has been encouraging recent growth of available genomic information in the Lathyrus genus these resources are still modest when compared with other legume crops such as pea As mentioned before in this review several neutral DNA marker systems have been applied successfully in Lathyrus diversity studies However this success has not been translated into gene discovery or development of trait associated markers for marker assisted selection MAS in Lathyrus breeding There is one report of an L sativus molecular marker linkage map developed to identify genomic regions linked to agronomically important traits ascochyta blight resistance Skiba et al 2004 Also there is no subsequent report on the use of the detected associated markers in breeding for resistance in Lathyrus Moreover this linkage map was not adequately saturated with markers presenting numerous gaps and short linkage groups Vaz Patto et al 2006 b due to the lack of anchor markers it could not be aligned and compared with other legume species linkage maps Earlier studies indicated that extensive genome conservation based on comparative genetic mapping was exhibited by members of the legume Papilionoideae subfamily such as Pisum Lens Vicia or Cicer Zhu et al 2005 There is an urgent need to develop a more comprehensive genetic map for Lathyrus with localization of useful genes and quantitative trait loci QTLs for MAS and with the possibility of alignment with other species in a comparative mapping approach The inclusion of cross amplified anchor markers needs to be addressed to allow comparative mapping with other related legume species opening the way for using Lathyrus as a source of interesting traits for other related species and vice versa Genomic and EST microsatellites were the most commonly attempted cross species amplification marker systems in Lathyrus but ITAP RGA and DR genes have been used on these cross amplification studies involving Lathyrus sp see above Some of these marker systems like microsatellites have an additional advantage for linkage map development since they are co dominant markers The incorporation of co dominant markers will be very important for a correct estimation of genetic distances among markers in repulsion phase Vaz Patto et al 2011 As previously mentioned above not only cross amplifiable markers from other legume species are being used in Lathyrus genetic studies Lathyrus EST are being made available in public databases in particular for L sativus and L odoratus and these are now being used to develop molecular markers associated with coding DNA Very recently cDNA libraries have also been developed for L cicera and EST SSR markers identified Almeida et al 2011 This marker system generally has a high degree of sequence conservation and may potentially be more transferable among species thus facilitating comparative genomic mapping Vaz Patto et al 2006 b With the development of high throughput and dense genotyping the assessment of the correlation between phenotype and genotype needed for the development of MAS approaches has shifted from focusing on two parental lines differing strongly in phenotype to populations of unrelated individuals Association mapping panels by sampling more genetic diversity can take advantage of many more generations of recombination and avoid the time consuming generations of crosses Morrell et al 2012 High throughput genotyping associated with a core collection evaluation will facilitate trait dissection and gene discovery through association mapping as well as characterization of genetic structure Cobb et al 2013 That is why it would be so important to concentrate the evaluation efforts on to a core sub set collection representative of all the existing diversity but of a manageable size Other biotechnological advances In contrast to the development of and now initiated use of different molecular markers with breeding objectives other biotechnological advances conceived particularly for functional studies are not currently used on Lathyrus breeding Expression analysis studies were initially performed on L sativus inoculated with Mycosphaerella pinodes using a limited number of 29 ESTs representing genes coding for enzymes and proteins involved in different levels of defence Skiba et al 2005 Some of the Lathyrus EST libraries previously mentioned in this review are now being developed not only for identifying molecular markers useful for the construction of high density genetic linkage maps but also for allowing expression analysis studies in order to identify and assess the function of putative genes thought to be involved in plant disease resistance responses This is the case of rust resistance in L sativus and L cicera Almeida et al 2012 Next generation sequencing technologies have also been recently applied to the L sativus Ascochyta sp interaction through SuperSAGE quantitative gene expression profiling Almeida et al 2013 With this approach it was possible to identify 3000 P 0 05 overexpressed or 900 P 0 05 underexpressed transcripts during the first 24 h after inoculation between infected and control tissue opening the way to a powerful route of identification of candidate resistance genes and the study of resistance gene networks in L sativus Gurung and Pang 2011 have recently prioritized the future construction of Lathyrus EST libraries from developing pods and seeds to achieve representation of reproductive tissues In addition these authors have called attention to the difficulties of proof of function studies of putative Lathyrus genes via overexpression deletion or silencing due to the non existence of a widely employed transformation system only one event reported by Barik et al 2005 or a broadly applicable virus induced gene silencing VIGS reverse genetics approach until now only reported for L odoratus Grønlung et al 2008 Likewise the reduced size of the presently available mutant populations does not provide the opportunity to study the effects of gene deletions silencing through targeting induced local lesions in genome TILLING approaches Gurung and Pang 2011 Mutation breeding has been employed on several occasions to create additional variability in a range of traits from plant growth habit to ODAP or methionine and lysine content reviewed in Vaz Patto et al 2011 Extensive variation was also detected in the course of tissue culture studies in several morphological traits and has also been exploited in grass pea breeding Kumar et al 2011 Embryo rescue and protoplast fusion protocols meant to increase the range of species in successful interspecific crosses Ochatt et al 2007 have unfortunately not been routinely used for grass pea improvement to date due to the difficult regeneration of hybrid plants There are also survival problems with the recovered confirmed haploid plants that were reported in the haplo diploidization method established from cultured isolated microspores in L sativus Ochatt et al 2009 hindering its application in breeding An in vitro protocol for fast production of advanced progeny that drastically shortens generation cycles has been developed in L sativus Ochatt et al 2004 Over three generations per year can be obtained instead of the normal two allowing a faster progress in L sativus improvement However this approach is only applicable when few seeds plant are intended as in single seed descendant breeding schemes Previous Section Next Section CONCLUSIONS Today due to the deluge of low cost genomic information phenotyping is quickly emerging as the major operational bottleneck and funding constraint of genetic analysis Cobb et al 2013 Consequently after overcoming the problem of availability of appropriate germplasm resources to address specific questions through the establishment of a core collection further emphasis should be placed on overcoming the shortage of high quality phenotypic information to associate with the high throughput genotyping information We propose that future international efforts on L sativus and L cicera improvement should concentrate on the development of publicly available joint core collections and on its high resolution genotyping This will be critical for permitting a decentralized phenotyping where multiple researchers can interrogate the same genetic materials phenotyping in environments and with technology and analytical expertise that are uniquely available to different research groups Cobb et al 2013 Such co ordinated international effort is sure to translate into more efficient and faster breeding approaches which are especially needed for such neglected but promising underutilized species Previous Section Next Section ACKNOWLEDGEMENTS The authors would like to acknowledge K P Colleary Ed D for English revision of the manuscript and the financial support of Fundação para a Ciência e a Tecnologia Portugal through grants PEst OE EQB LA0004 2011 and PTDC AGR GPL 103285 2008 and Research Contract by the Ciência 2008 program to M C V P This research was also funded by FP7 ARIMNet MEDILEG project and by Spanish project AGL2011 22524 Sponsorship by the Annals of Botany to D R to give a key lecture at the 1st Legume Conference at Novi Sad Serbia is also acknowledged The Author 2014 Published by Oxford University Press on behalf of the Annals of Botany Company All rights reserved For Permissions please email journals permissions oup com Previous Section LITERATURE CITED Abd El Moneim AM van Dorrestein B Baum M Ryan J Vejiga G Role of ICARDA in improving the nutritional quality and yield potential of grasspea Lathyrus sativus L for subsistence farmers in dry areas Lathyrus Lathyrism Newsletter 2001 2 55 58 Google Scholar Abd El Moneim AM Nakkoul H Masri S Ryan J Implications of zinc fertilization for ameliorating toxicity neurotoxin in grasspea Lathyrus sativus Journal of Agricultural Science and Technology 2010 12 69 78 Web of Science Google Scholar Ali HBM Meister A Schubert I DNA content rDNA loci and DAPI bands reflect the phylogenetic distance between Lathyrus species Genome 2000 43 1027 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