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Afila, a new mutant in pea (Pisum sativum L.)
... The af or semi-leafless phenotype in pea arises as a consequence of a recessive mutation at a single locus (Kujala, 1953;Solov'eva, 1958;Goldenberg, 1965;Khangildin, 1966;Swiecicki, 1982) on linkage group (LG) I (Marx, 1969;Blixt, 1972;Bordat et al., 2011;Tayeh et al., 2015a). In WT (AfAf ) genotypes, the proximally positioned lateral primordia, on the flanks of the leaf primordium, develop into determinate leaflet laminae, but in af mutants, this fails to happen; the lateral and apical primordia behave in the same way so that second-order, midrib-like, primordia in turn initiate organ primordia (Meicenheimer et al., 1983;Côté et al., 1992). ...
... Besides the lack of information on the molecular identity of af, it was also unclear how the pea community managed to transform af from simple spontaneous mutations observed in the USSR (Solov'eva, 1958), Finland (Kujala, 1953) and Argentina (Goldenberg, 1965) to a major breeding criterion in dry pea. Did pioneer breeders simply rely on spontaneous mutations in their programmes? ...
... Much of the yield was lost at harvest due to lodging, and seed quality was often poor due to pathogen attacks arising from damp soil conditions (Banniza et al., 2005). The success story of improving lodging resistance in pea owes much to the introduction of the af mutation (Kujala, 1953;Solov'eva, 1958;Goldenberg, 1965;Khangildin, 1966;Swiecicki, 1982) together with a mutation in Mendel's stem length gene Le (White, 1917;Lester et al., 1997;Martin et al., 1997) into breeding material. The af mutation results in leaves with an increased number of tendrils, whereas the le mutation causes short-stem internodes (Lester et al., 1997;Martin et al., 1997). ...
The afila (af) mutation causes the replacement of leaflets by a branched mass of tendrils in the compound leaves of pea – Pisum sativum L. This mutation was first described in 1953, and several reports of spontaneous af mutations and induced mutants with a similar phenotype exist. Despite widespread introgression into breeding material, the nature of af and the origin of the alleles used remain unknown.
Here, we combine comparative genomics with reverse genetic approaches to elucidate the genetic determinants of af. We also investigate haplotype diversity using a set of AfAf and afaf cultivars and breeding lines and molecular markers linked to seven consecutive genes.
Our results show that deletion of two tandemly arranged genes encoding Q‐type Cys(2)His(2) zinc finger transcription factors, PsPALM1a and PsPALM1b, is responsible for the af phenotype in pea. Eight haplotypes were identified in the af‐harbouring genomic region on chromosome 2. These haplotypes differ in the size of the deletion, covering more or less genes.
Diversity at the af locus is valuable for crop improvement and sheds light on the history of pea breeding for improved standing ability. The results will be used to understand the function of PsPALM1a/b and to transfer the knowledge for innovation in related crops.
... The principal approach being pursued for the genetic dissection of processes that control leaf formation in P. sativum consists of characterization of relative morphology, anatomy and gene expression in leaves at different stages of development, in leaf morphology affecting recessive loss-of-function mutants and corresponding wildtypes [18,20,22,23,[28][29][30]40]. Concordantly, spontaneous and leaf morphology mutants have been detected/are being searched in land races, segregating populations of wide crosses and mutagenized populations [5,[11][12][13][14][15][16][17]19,21,24,27,36,38,40]. The comparable plants of the genotypes harboring such mutations singly and in various permutations and combinations, in homozygous and/or heterozygotes conditions, are subjected to detailed examination [25,[30][31][32][33][34][35][37][38][39][40]. ...
... The characteristics of mutations in the genes UNIFOLI-ATA (UNI) or UNI-TENDRILLED ACCACIA (UNI-TAC), TENDRILL-LESS (TL) and AFILA (AF) have provided the following view on the developmental processes that determine fates of regional domains along the acropetal long axis of leaf-blade in P. sativum [1,4,[11][12][13][14][15][16][17][18][19][20][21][24][25][26][27][28][29][30][31]34,39,40]. (a) UNI gene activity is required for the full growth of rachis or elaboration of all three domains of rachis in leaf-blade. ...
... The line SKP-100 was also crossed with three other lines: SKP-2, a line bearing tl/tl acacia-type leaves, SKP-3, a line bearing af/af specified tendrils in place of leaflets and SKP-4, a double mutant line bearing tl/tl, af/af genotype in which compound pinnae are formed in proximal and distal leaf-blade domains. The af and tl alleles used have been previously described [11,12,15,16]. Plants bearing af/af, mfp/mfp; tl/tl mfp/mfp; and af/af, tl/tl, mfp/mfp were recovered in F 2 generation on the basis of distinct morphologies of their leaves and tested for homozygosity at the concerned loci by raising their M 3 generation. ...
The wild type compound leaf-blade of Pisum sativum has one to three pairs of simple leaflet pinnae in its petiole proximal domain, one to four pairs of simple tendril pinnae in the distal domain and a simple tendril pinna in the apical domain. A novel ethyl methane sulfonate induced dominant mutant was isolated and characterized whose leaf-blades formed MULTIFOLIATE PINNA pairs in the distal domain. The distal multifoliate pinnae or compound pinna-blades had three tendrilled-leaflets as pinnules. The pinnules had a bifacial elliptic-lanceolate leaflet body and arc shaped apex that mimicked the ringlet shaped apex of tendrils. The TL/tl, mfp/mfp and tl/tl, mfp/mfp leaf-blades also produced multifoliate (compound) pinna-blades in distal positions; the pinnules of these genotypes had elliptic shape. The pinnae were branched tendrils in TL/TL, MFP/mfp plants. The leaf-blade rachis was more ramified in af mfp double mutants than in af mutant. In the af mfp double mutant, the multifoliate pinna-blades were present on tertiary and secondary branches of the rachis in the proximal domain and on secondary branches and the primary rachis in the distal domain. The leaf-blades of the af tl mfp triple mutant genotype were an order of magnitude more ramified than those of af tl and af mfp genotypes in proximal as well as distal domains. The leaf-blade phenotypes of various genotypes revealed in this study and those known from previous work have allowed the following conclusions about the nature of mfp mutation and mfp function(s). (a) The presence of mfp mutation or mfp function changes the identity of distal primordia, from tendrils in the wildtype (MFP/MFP) leaf-blades to multifoliate pinna-blades in mfp/mfp mutant. (b) A pathway for the lamination of pinnules of multifoliate blades formed in distal and terminal domains in the mfp mutant and all domains in af mfp double mutant is activated by the mfp mutation. (c) The leaflet-/pinnule-lamination pathway activated by the tl mutation interacts with the mfp-directed pathway. (d) The mfp mutation intensifies rachis ramification in proximal and distal domains activated by the af mutation. This process is distinct from analogous rachis ramification that occurs in the af tl double mutant.
... One consequence of the af mutation (ICujala, 1953;Goldenberg, 1965) is that lateral leaflet primordia are converted into lateral rachis primordia (Gould etal., 1986;Cote et al., 1992;Gourlay et al., 2000); the latter themselves have the potential to initiate further lateral primordia. The af leaf, when combined with the // mutation, develops as a supercompound leaf with even more ramified raches each terminating in miniature leaflets (Goldenberg, 1965;Gould et al., 1992) (Fig. 7A). ...
... One consequence of the af mutation (ICujala, 1953;Goldenberg, 1965) is that lateral leaflet primordia are converted into lateral rachis primordia (Gould etal., 1986;Cote et al., 1992;Gourlay et al., 2000); the latter themselves have the potential to initiate further lateral primordia. The af leaf, when combined with the // mutation, develops as a supercompound leaf with even more ramified raches each terminating in miniature leaflets (Goldenberg, 1965;Gould et al., 1992) (Fig. 7A). Marx (1987) emphasized the threedimensional aspect of af and aftl leaves, whereas twodimensional diagrams represented them as bipinnate, Arrow indicates a node with an axillary inflorescence bearing a mutant flower, subtended by a unifoliate leaf. ...
The partial-shoot theory of the leaf was a controversial hypothesis revived by Arber and supported by her morphological and
anatomical studies. This theory highlighted the parallels between leaves and shoots and contrasted with an alternative view
that leaves, with their limited growth potential, are completely distinct from shoots. Pea morphological mutants with altered
growth potential in their compound leaves are described. The unifoliata mutant has a limited growth potential relative to wild-type;cochleata, afila and insecatus have extended potentials. Characterization of theunifoliata mutation and gene expression patterns indicate that unifoliata is a common factor in pea compound leaf and floral shoot development, and so provides rudimentary support for the idea that
some leaves have shoot-like characteristics. Tomato leaves are also considered to lend tentative support. The afila and insecatus leaf forms are described as bipinnate and weakly bipinnate, respectively. These and the tendril-less mutant are potential phenocopies of legume relatives, an idea based on Vavilov's law of homologous series of variation. Arber
illustrated, but did not articulate in genetical terms, that morphological variation in structures within an individual plant
can be interpreted as reiteration of design. Analogous with Vavilov's view, this can be considered a consequence of the same
genetic programme in a different location.Copyright 2001 Annals of Botany Company
... Svoboda. Later on Goldenberg (1965) described afila as spontaneous mutation from the cv. Cuareton. ...
Pesic, V., R. DjoRDjeVic, e. KADHUM, P. jAnKoVic and D. Misic, 2013. Influence of the afila gene on grain yield in pea (Pisum sativum L.). Bulg. J. Agric. Sci., 19: 186-193 The aim was to investigate the possibility of introducing leafless or semi-leafless features from afila cultivar "Filigreen" in early, mid-early and late pea cultivars. F 1 and F 2 generations were grown together in a single nursery during year 2006. F 1 plants with normal leaves were obtained thus proving that afila is a recessive characteristic. The total of 1490 F 2 plants segregated with 2.93 plants with normal leaves to 1.07 with afila leaves. Chi-square test confirmed monohybrid model of inheritance 3:1. Afila plants with differing maturity dates were obtained. The best recombination's, in terms of grain yield per plant, were obtained by crossing "Filigreen" x "Maja" (early cultivar), "Filigreen" x "Oskar" (mid-early cultivar) and "Filigreen" x "Zelena dolina" (late cultivar). Grain yield of F 2 afila plants was reduced by 6-18% in relation to grain yield of normal plants (depending on parents, combinations). Grain yield was reduced, in average, by 10% for afila plants compared to normal (leafy) plants in the F 2 for all possible crosses.
... The leaf type could be conventional, semileafless or leafless (Davies et al. 1985). Goldenberg (1965) found -as a result of spontaneous mutation -the afila plant type whose leaflets converted to tendrils. Jaranowski (1976) obtained a similar plant type via induced mutagenesis and started to use this type in breeding programmes. ...
This review recapitulates the history, important milestones, the current status, and the perspectives of the pea (Pisum sativum L.) transformation as a tool for pea crop breeding. It summarises the developments of the pea transformation from the first methodological experiments to achieving the complete transformation and regeneration of genetically modified (GM) plants, transformation with the first genes of interest (GOI), to recent techniques of targeted genome editing. We show how recent biotechnological methods and genetic engineering may contribute to pea breeding in order to speed up the breeding process and for the creation of new pea cultivars. The focus is laid on genetic engineering which represents an excellent technology to enhance the pea gene pool with genes of interest which are not naturally present in the pea genome. Different methods of pea transformation are mentioned, as well as various GOI that have been used for pea transformation to date, all aimed at improving transgenic pea traits. Tolerance to herbicides or resistance to viruses, fungal pathogens, and insect pests belong, among others, to the pea traits that have already been modulated by methods of genetic engineering. The production of phytopharmaceuticals is also an important chapter in the use of genetically modified peas. We compare different methods of introducing transgenes to peas and also the usage of different selective and reporter genes. The transformation of other major legumes (soybeans, beans) is marginally mentioned. The effect of genetically modified (GM) peas on animal health (feeding tests, allergenicity) is summarised, the potential risks and benefits of pea modification are evaluated and also the prime expectations of GM peas and the real current state of this technology are compared. Unfortunately, this technology or, more precisely, the products created by this technology are under strict (albeit not scientifically-based) legislative control in the European Union.
... The replacement of the tendrils in the leaf of wild type by leaflets is controlled by a single recessive gene tendril (tl) and creates "acacia" phenotype (de Vilmorin and Bateson 1911). On the contrary, the single recessive gene afila (af) converted leaflets into tendrils (Kujala 1953;Goldenberg 1965). The two recessive genes af tl altered the identity of leaf pinnae, afila (af), and acacia (tl) DeMason 1997, 2000). ...
Field pea (Pisum sativus L.) is a nutritionally dense winter season pulse crop, consumed worldwide as food, feed and fodder and offers nutritional security to low-income folks of various developing countries. It is an excellent source of protein and carbohydrate in juxtaposition with vitamins, essential amino acids, and macro- and micronutrients. In addition, it plays an important role in management of Type 2 diabetes and body weight, blood cholesterol reduction, improves cardiovascular health and gastrointestinal function. It is susceptible to many biotic and abiotic stresses that seriously hinder its sustainable production. Over the years, sincere efforts have been made toward the genetic improvement of field pea to subsidize antinutritional components and elevate production potential. In this book chapter, the importance of the crop, its common uses, origin, evolution, gene pool, botanical description, floral biology, cytogenetics and molecular cytogenetics, genetic variability for important agronomic traits, inheritance of qualitative and quantitative traits, and brief account of genetic resources have been illustrated. The achievement made in field pea through conventional and nonconventional breeding approaches, that is, hybridization, distant hybridization, and mutation breeding, have been reviewed. The current scenario of genomics resources and marker-assisted breeding has also been deliberated. Moreover, the breeding objectives, major constraints, and future perspectives in order to explore cutting-edge tools and technique for enriching field pea genomic resources have been outlined. Furthermore, the currently existing coordinated testing system for new entries and quality seed production has also been described in short. Overall, to accelerate genetic gain in field pea along with nutritional enrichment, there is urgent need of exploitation of recent advanced tools and techniques such as transcriptomics, proteomics, metabolomics, small RNAomics, epigenomics, interactomics, bioinformatics, genomic selection, genome editing, and speed breeding to bolster the field pea breeding program.Keywords
Pisum sativum
CytogeneticsGermplasmBreeding strategiesBreeding objectivesCoordinated system of testing
... Dada la condición afila de las líneas evaluadas producida por un gen mutante que remplaza hojas por zarcillos (Goldenberg 1965), la cual contrasta con los testigos de hojas normales, y considerando que los padres donantes del gen afila son plantas arbustivas y los padres recurrentes usados en los retrocruzamientos son variedades volubles, el objetivo inicial de los cruzamientos realizados se ha cumplido al encontrar líneas que teniendo dicho gen (afila), producen rendimientos similares a variedades comerciales en este caso Andina y Sindamanoy. Además estas líneas presentan la reacción entre resistente y moderadamente resistente al patógeno Erysiphe polygoni. ...
In the past five years in the department of Nariño, the increase in severity of damage produced by oidio (Erysiphe polygoni) in vetch has been observed, which produces loss of yield crop between 20 and 40%. The improved varieties present susceptibility to the pathogen. This research was conducted in SENA's Lopez Farm in Pasto-Colombia, in two phases. In the first phase, 90 lines of pea were evaluated (Pisum sativum L.) along with the gen afila for yield and reaction to powdery mildew (Erysiphe polygoni) under natural inoculate conditions. The assessed lines were obtained by simple crosses and backcrosses between the parentals Andina, San Isidro, and Sindamanoy, and the donor genotypes of the afila gen: Dove, ILS3568 and ILS3575. A random augmented blocks design was used. Results showed that 10% of the materials were resistant to powdery mildew, 20% were moderately resistant, 60% were moderately susceptible, and 10% were susceptible. From these materials, 19 resistant and moderately resistant genotypes with the highest yield were selected, which were assessed in the second phase for yield and components. For green pod yield and dry grain, 84% and 68.4% of the tested lines reached similar averages than those in the commercial controls Sindamanoy and Andina. The selected lines with the gen afila recovered 88,42% of the yield and yield components from their parents Andina, San Isidro, and Sindamanoy.
... However, the differences for LF expression within groups indicate that LF transcript level is not sufficient to explain the node of flowering. (Goldenberg, 1965)), hence decreasing the leaf volumes proportionally to the meristem and artificially increasing LF transcripts levels in af genotypes compared to Af genotypes. Analysis of Af/af isolines should allow confirmation of AF effect on LF expression. ...
Le pois (Pisum sativum), par son double statut d’espèce modèle pour l’étude du développement, et d’espèce agronomique, représente une espèce modèle idéale pour des études intégrées à différentes échelles biologiques. La transition florale est un caractère clé du développement et des approches variées ont conduit à l’obtention de nombreuses données pour la floraison chez le pois : (i) les approches de génétique et physiologie menées en conditions contrôlées sur une large gamme de mutants ont conduit au développement d’un modèle descriptif, mais sans capacité de prédiction, développant les interactions entre les gènes connus contrôlant la floraison ; (ii) l’étude approfondie en conditions de plein champ du contrôle de la floraison a permis de développer des modèles écophysiologiques de la date de floraison en fonction de la photopériode et de la température à forte capacité de prédiction mais qui ne prennent pas en compte le génotype. Plus récemment, les données sur Arabidopsis thaliana permettent d’avoir une compréhension au niveau moléculaire des mécanismes en jeu. Ce projet est une première approche pour intégrer ce large jeu de données au sein d’un modèle prédictif de la date d’initiation florale, décomposé sous la forme du produit mathématique du premier noeud d’initiation florale (NFI) et du temps nécessaire à l’initiation d’un nouveau noeud à l’apex (plastochrone). Un premier modèle mathématique pour la régulation génétique du NFI a été développé qui permet de prédire le NFI pour différents génotypes et photopériodes. Les réponses du NFI et du plastochrone aux conditions environnementales et en particulier à la photopériode ont été analysées précisément. Afin de compléter le modèle, je me suis intéressée particulièrement aux deux gènes clés de floraison, LATE FLOWERING (LF) et HIGH RESPONSE TO PHOTOPERIOD (HR). Des approches moléculaires pour HR ont permis de montrer que le gène était impliqué dans la voie de transduction de la lumière vers l’horloge circadienne, et de nouveaux gènes candidats ont été proposés. Ce travail propose des pistes pour exploiter l’approche de modélisation pour la floraison chez le pois à la lumière des nouvelles données moléculaires.
... It was dwarfing genes in wheat and rice that allowed the development of varieties that powered the Green Revolution (Hedden, 2003). In pea breeding a breakthrough in lodging resistance came in the 1980's with the development of dwarf (le) varieties with the semi-leafless (afila) trait (Goldenberg, 1965). Le, also known as Mendel's tall/ dwarf gene (Mendel, 1866), influences plant height and stem diameter (Smitchger and Weeden, 2019), with the recessive le allele causing the dwarfing phenotype. ...
... WAX2 is among the genes controlling wax biosynthesis in Arabidopsis [47], and glossy13 is another gene with similar role reported in maize [48]. Lodging contributes to canopy heating in pea, and upright and semileafless cultivars with the afila gene have cooler CT [12,49]. Tar'an et al. [50] identified major loci associated with lodging resistance in pea on LG III, and one of the markers we identified for CT is also on LG III, suggesting genes controlling lodging also control CT. ...
Environmental stress hampers pea productivity. To understand the genetic basis of heat resistance, a genome-wide association study (GWAS) was conducted on six stress responsive traits of physiological and agronomic importance in pea, with an objective to identify the genetic loci associated with these traits. One hundred and thirty-five genetically diverse pea accessions from major pea growing areas of the world were phenotyped in field trials across five environments, under generally ambient (control) and heat stress conditions. Statistical analysis of phenotype indicated significant effects of genotype (G), environment (E), and G × E interaction for all traits. A total of 16,877 known high-quality SNPs were used for association analysis to determine marker-trait associations (MTA). We identified 32 MTAs that were consistent in at least three environments for association with the traits of stress resistance: six for chlorophyll concentration measured by a soil plant analysis development meter; two each for photochemical reflectance index and canopy temperature; seven for reproductive stem length; six for internode length; and nine for pod number. Forty-eight candidate genes were identified within 15 kb distance of these markers. The identified markers and candidate genes have potential for marker-assisted selection towards the development of heat resistant pea cultivars.
... Other genes are involved in the leaf primordium organogenesis and in the determination of the compound leaf architecture. Depending on the leaf portions, the negative regulation of PsUNI expression by PsTENDRIL-LESS (PsTL) promotes the distal and terminal tendrils identity (de Vilmorin and Bateson, 1911), PsUNI repression by PsAFILA (PsAF) gene promotes the proximal leaflet identity (Kujala, 1953;Goldenberg, 1965) and PsUNI repression by PsCOCHLEATA1 (PsCOCH1) gene confers the stipule identity (Wellensiek, 1959;Blixt, 1967;Gourlay et al., 2000;Kumar et al., 2009;Couzigou et al., 2012). The Pscoch1 mutant displays a wide range of stipule phenotypes that can be explain by different reasons: i) The CHAPTER II. ...
The symbiotic interaction between legumes andrhizobia results in the formation of a symbiotic nitrogen fixingnodule.This de-novo generated symbiotic organ allows the intracellularaccommodation of the rhizobia which reduces through theirnitrogenase activity the atmospheric nitrogen in ammonium, anitrogen form usable by the host plant.The molecular mechanisms underlying the symbiotic partnersrecognition, the infection process and the nodule organogenesis arewell described, however the identity establishment and maintenanceof this unique underground organ remain mis-understood.The Medicago truncatula NODULE-ROOT, the Arabidopsisthaliana BLADE-ON-PETIOLE and the Pisum sativumCOCHLEATA genes are members of a highly conserved NOOTBOP-COCH-LIKE1 (NBCL1) specific clade that belongs to theNON-EXPRESSOR OF PATHOGENESIS RELATED PROTEIN1-LIKE gene family. In legumes, the members of this NBCL1 cladeare known as key regulators of the symbiotic nodule identity.The present thesis work aims to better understand the roles of theNBCL1 genes, in both indeterminate and determinate nodule formingspecies and to discover new molecular actors involved in theNBCL1-dependent regulation of the nodule identity essentially usingnovel TILLING, Tnt1 and LORE1 insertional mutants in three legumespecies, Medicago, Pisum and Lotus.This thesis work has allowed the identification and thecharacterization of new mutants for genes belonging to a secondarylegume-specific NBCL2 sub-clade. We revealed that the members ofthis legume-specific NBCL2 sub-clade play important roles in noduledevelopment, identity establishment and maintenance, andconsequently in the success and in the efficiency of the symbioticassociation.This thesis work also shows that during evolution, the noduledevelopmental program has recruited pre-existing regulatorymechanisms for the nodule development and identity, such as theregulatory module involving interactions between NBCL proteins andTGACG type basic leucine zipper transcription factors. We identifiedthe transcription factor, MtPERIANTHIA-LIKE, as a first interactingpartner of NBCL proteins in a context of root nodule symbiosis.NBCL genes are also involved in the regulatory networks thatcontrol the development and the determinacy of many abovegroundvegetative and reproductive organs and were also shown as involvedin their abscission ability.In this thesis we also explored the roles of these highly conservedNBCL genes in the development of the non-domesticated grass,Brachypodium distachyon.
... Plant structure is often modified in crop plants. In legumes, leaf architecture is regulated by a series of genes (Gourlay et al. 2000), but the most useful in the development of the pea crop was afila (Goldenberg 1965), the so-called semi-leafless character. An afila mutant showing leaflets converted to tendrils was incorporated into the pea breeding program at the John Innes Centre in the 1980s (Snoad 1981), its advantage being that the plants remain more erect because of the support each gives to the other through the enhanced number of tendrils. ...
Grass pea (Lathyrus sativus) is a hardy legume grown for food, feed and fodder. It is an ancient crop, having been cultivated for more than 8000 years because of its tolerance of drought, flooding, salinity and poor soils, its ability to fix nitrogen and its seeds with high levels of protein. These traits make it an outstanding crop for ensuring nutritional security (particularly for protein) for resource-poor farmers, especially in the face of impending changes in climate. However, the presence of β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP or ODAP), a neurotoxin present in the seeds and vegetative tissues of grass pea, has limited its breeding and modern-day cultivation. β-ODAP causes lathyrism, a paralysis of the lower limbs that occurs in epidemics in under-nourished communities. This has resulted in grass pea being an “orphan crop” whose potential has not been fully realized due to lack of markets and research funding. The recent emphasis on climate smart crops has refocused attention on this very promising crop. Genomic resources and low ODAP lines are being developed, and it is hoped that these will soon allow grass pea to reach its full potential as a resilient protein crop for food and nutritional security through sustainable agriculture in the face of climate change.
... Plant structure is often modified in crop plants. In legumes, leaf architecture is regulated by a series of genes (Gourlay et al. 2000), but the most useful in the development of the pea crop was afila (Goldenberg 1965), the so-called semi-leafless character. An afila mutant showing leaflets converted to tendrils was incorporated into the pea breeding program at the John Innes Centre in the 1980s (Snoad 1981), its advantage being that the plants remain more erect because of the support each gives to the other through the enhanced number of tendrils. ...
Grass pea (Lathyrus sativus) is a hardy legume grown for food, feed, and fodder. It is an ancient crop which has been cultivated for more than 8000 years because of its tolerance of drought, flooding, salinity, and poor soils, its ability to fix nitrogen, and its seeds with high levels of protein. These traits make it an outstanding crop for ensuring nutritional security (particularly for protein) for resource-poor farmers, especially in the face of impending changes in climate. However, the presence of β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP or ODAP), a neurotoxin present in the seeds and vegetative tissues of grass pea, has limited its breeding and modern-day cultivation. β-ODAP causes lathyrism, a paralysis of the lower limbs that occurs in epidemics in undernourished communities. This has resulted in grass pea being an “orphan crop” whose potential has not been fully realized due to lack of markets and research funding. The recent emphasis on climate smart crops has refocused attention on this very promising crop. Genomic resources and low-ODAP lines are being developed, and it is hoped that these will soon allow grass pea to reach its full potential as a resilient protein crop for food and nutritional security through sustainable agriculture in the face of climate change.
... Two dwarf isogenic lines of "New Line Early Perfection" peas were used in these experiments: the mutants acacia and parsley-leaved (Marx, 1974Marx, , 1987Snoad, 1974). These two varieties differ only at the afila (aj) locus (Kujala, 1953;Goldenberg, 1965;Gottschalk, 1972;Marx, 1974;Snoad, 1974) and were recessive at both the stipule (st) and tendrilled (tl) loci (acacia peas: Af/-; stlst; tlltl, parsley-leaved peas: aflaf; stlst; tlltl). Although differing at only a single locus, these two genotypes have very different forms. ...
We investigated the hypothesis that plant form can dramatically affect plant competitive ability, and that forms with dense canopies can invade populations of plants with more open canopies regardless of initial relative frequencies. Under controlled field conditions, we examined the effects of plant form on growth rate, size variation, mortality, and reproduction in high-density monocultures and mixtures of two morphologically distinct varieties of peas. These two varieties differ genetically at only the afila locus. In high-density monocultures and mixtures, peas with finely dissected, minute leaflets (af/af) grew more slowly and produced fewer seeds than Af/-individuals with large leaflets that cast more shade on neighbors. After as few as four generations, mixtures begun with 10% Af/- peas would be expected to evolve to Af/- monocultures. We conclude that an increase in morphological complexity (e.g., virtually leafless to leafy) can have dramatic ecological and evolutionary impacts on plant population dynamics.
... Dada la condición afila de las líneas evaluadas producida por un gen mutante que remplaza hojas por zarcillos (Goldenberg 1965), la cual contrasta con los testigos de hojas normales, y considerando que los padres donantes del gen afila son plantas arbustivas y los padres recurrentes usados en los retrocruzamientos son variedades volubles, el objetivo inicial de los cruzamientos realizados se ha cumplido al encontrar líneas que teniendo dicho gen (afila), producen rendimientos similares a variedades comerciales en este caso Andina y Sindamanoy. Además estas líneas presentan la reacción entre resistente y moderadamente resistente al patógeno Erysiphe polygoni. ...
En los últimos cinco años en el departamento de Nariño se observa incremento de la severidad del ataque de oídio (Erysiphe polygoni) en arveja que produce pérdidas en rendimiento entre un 20 y un 40%. Las variedades mejoradas presentan susceptibilidad al patógeno. La investigación se realizó en la granja Lope del SENA, Pasto, Colombia. En la primera fase, se evaluaron 90 materiales de arveja (Pisum sativum) con el gen afila por rendimiento y reacción al oídio (Erysiphe polygoni) bajo condiciones de inoculo natural. Las líneas evaluadas se obtuvieron por cruzamientos simples y retrocruzamientos entre los parentales Andina, San Isidro y Sindamanoy y los genotipos donantes del gen afia: Dove, ILS3568 e ILS3575. Se utilizó un diseño de bloques al azar aumentado. Se encontró que 10% de los materiales fueron resistentes a oidio, 20% moderadamente resistentes, 60% moderadamente susceptibles y 10% susceptibles. De estos materiales se seleccionaron 19 genotipos resistentes, y moderadamente resistentes y de mayor rendimiento, los cuales fueron evaluados en la segunda fase por rendimiento y sus componentes. Para rendimiento en vaina verde y en grano seco el 84% y el 68,4% de las líneas evaluadas tuvieron promedios similares a los testigos comerciales Andina y Sindamanoy. Las líneas seleccionadas con el gen afila recuperaron en un 88,42% las características de rendimiento y sus componentes de los progenitores Andina, San Isidro y Sindamanoy
... Svoboda. Later on Goldenberg (1965) described afila as spontaneous mutation from the cv. Cuareton. ...
The aim was to investigate the possibility of introducing leafless or semi-leafless features from afila cultivar "Filigreen" in early, mid-early and late pea cultivars. F1 and F2 generations were grown together in a single nursery during year 2006. F1 plants with normal leaves were obtained thus proving that afila is a recessive characteristic. The total of 1490 F2 plants segregated with 2.93 plants with normal leaves to 1.07 with afila leaves. Chi-square test confirmed monohybrid model of inheritance 3:1. Afila plants with differing maturity dates were obtained. The best recombination's, in terms of grain yield per plant, were obtained by crossing "Filigreen" x "Maja" (early cultivar), "Filigreen" x "Oskar" (mid-early cultivar) and "Filigreen" x "Zelena dolina" (late cultivar). Grain yield of F2 afila plants was reduced by 6-18% in relation to grain yield of normal plants (depending on parents, combinations). Grain yield was reduced, in average, by 10% for afila plants compared to normal (leafy) plants in the F2 for all possible crosses.
... Yield gains of approximately 2% per year have been achieved . Lodging resistance has been improved through deployment of the afila gene for semi-leafless type (Kujala, 1953;Goldenberg, 1965) and secondarily through selection for increased stem strength (Banniza et al., 2005). Powdery mildew resistance based on the single recessive gene er-1 (Harland, 1948) has been widely deployed. ...
Pea (Pisum sativum L.) is an annual cool-season legume and one of the oldest domesticated crops. Dry pea seeds contain 22–25% protein, complex starch and fiber constituents, and a rich array of vitamins, minerals, and phytochemicals which make them a valuable source for human consumption and livestock feed. Dry pea ranks third to common bean and chickpea as the most widely grown pulse in the world with more than 11 million tons produced in 2013. Pea breeding has achieved great success since the time of Mendel's experiments in the mid-1800s. However, several traits still require significant improvement for better yield stability in a larger growing area. Key breeding objectives in pea include improving biotic and abiotic stress resistance and enhancing yield components and seed quality. Taking advantage of the diversity present in the pea genepool, many mapping populations have been constructed in the last decades and efforts have been deployed to identify loci involved in the control of target traits and further introgress them into elite breeding materials. Pea now benefits from next-generation sequencing and high-throughput genotyping technologies that are paving the way for genome-wide association studies and genomic selection approaches. This review covers the significant development and deployment of genomic tools for pea breeding in recent years. Future prospects are discussed especially in light of current progress toward deciphering the pea genome.
... The single laminate form for which the mutation is named is shown in Figure 1B; however, leaves on a uni mutant plant may also be lobed, bi-, and trifoliate. The semidominant tl mutation (de Vilmorin and Bateson, 1911) replaces distal and terminal tendrils with leaflets ( Figure 1C), whereas the recessive af mutation (Kujala, 1953;Goldenberg, 1965) replaces leaflets with branching rachides bearing tendrils ( Figure 1D). In the af tl double mutant, branching rachides at proximal, distal, and terminal positions terminate in small leaflets ( Figure 1E). ...
The compound leaf primordium of pea represents a marginal blastozone that initiates organ primordia, in an acropetal manner, from its growing distal region. The UNIFOLIATA (UNI) gene is important in marginal blastozone maintenance because loss or reduction of its function results in uni mutant leaves of reduced complexity. In this study, we show that UNI is expressed in the leaf blastozone over the period in which organ primordia are initiated and is downregulated at the time of leaf primordium determination. Prolonged UNI expression was associated with increased blastozone activity in the complex leaves of afila (af), cochleata (coch), and afila tendril-less (af tl) mutant plants. Our analysis suggests that UNI expression is negatively regulated by COCH in stipule primordia, by AF in proximal leaflet primordia, and by AF and TL in distal and terminal tendril primordia. We propose that the control of UNI expression by AF, TL, and COCH is important in the regulation of blastozone activity and pattern formation in the compound leaf primordium of the pea.
... In a major review of pea leaf development mutants, Marx (1987) suggested that various mutants appear to operate in these three dierent`domains' within the leaf (basal, proximal and distal), and that the action of these leaf development genes might help to de®ne these areas. However, studies of pea leaf development have concentrated on the recessive mutant a®la(af) (Kujala, 1953;Goldenberg, 1965), the semidominant mutant tendril-less(tl) (Vilmorin and Bateson, 1911) and the af tl double mutant (Fig. 1D, F and G), which alter lea¯et and tendril pinna identity in the proximal and distal domains (Meicenheimer et al., 1983;Gould et al., 1986Gould et al., , 1994Cote et al., 1992;Lu et al., 1996;Villani and DeMason, 1997, 1999a, b, 2000. The stipules and basal compartment have received little attention, with only one model of pea leaf development (Gourlay et al., 2000) considering the cochleata (coch) mutant (Wellensiek, 1959), in which the stipules of the middle nodes of the plant are compound (Fig. 1B). ...
The stipule mutant cochleata(coch) and the simple-leaf mutantunifoliata (uni) are utilized to increase understanding of the control of compound leaf and flower development in pea. The phenotype of the coch mutant, which affects the basal stipules of the pea leaf, is described in detail. Mutant coch flowers have supernumerary organs, abnormal fusing of flower parts, mosaic organs and partial male and female sterility. The wild-type Coch gene is shown to have a role in inflorescence development, floral organ identity and in the positioning of leaf parts. Changes in meristem size may be related to changes in leaf morphology. In the coch mutant, stipule primordia are small and their development is retarded in comparison with that of the first leaflet primordia. The diameter of the shoot apical meristem of the uni mutant is approx. 25% less than that of its wild-type siblings. This is the first time that a significant difference in apical meristem size has been observed in a pea leaf mutant. Genetic controls in the basal part of the leaf are illustrated by interactions between coch and other mutants. The mutantcoch gene is shown to change stipules into a more ‘compound leaf-like’ identity which is not affected by thestipules reduced mutation. The interaction of coch and tendril-less(tl) genes reveals that the expression of the wild-type Tl gene is reduced at the base of the leaf, supporting the theories of gradients of gene action.
... The wildtype leaf blade in pea is compound and is composed of proximal pairs of leaflets and distal pairs of tendrils and typically terminates in a single tendril. The afila (af ) mutation alters the normal morphological pattern such that leaflets are replaced by compound (branched) tendrils (Kujala, 1953 ;Goldenberg, 1965 ;Lu et al., 1996), whereas the acacia\tendrilless (tl ) mutation causes distal tendrils to be replaced by leaflets (Vilmorin and Bateson, 1912 ;White, 1917). Leaves of aftl double mutant plants (phenotype known as pleiofila) possess compound (branched) pinnae that terminate in small leaflets at all positions (Marx, 1987 ;Villani and DeMason, 1997). ...
The wildtype leaf of the garden pea possesses proximal pairs of leaflets and distal pairs of tendrils in the blade region. Theafila (af) mutation causes leaflets to be replaced by compound (branched) tendrils. We characterized the morphological variation in leaf form along the plant axis and leaf development in early and late postembryonic leaves onafilaplants to infer the role of theAfgene. Leaf forms are more diverse early in shoot ontogeny onafilaplants.Afinfluences pinna length and pinna branching in addition to pinna type. Pinna initiation in the proximal region ofafilaleaf primordia is basipetal and delayed compared to wildtype plants. In addition, pinna development in the proximal region ofafilaleaves occurs for a longer period of time than on wildtype leaf primordia. Therefore,Afregulates the timing and direction of leaf developmental processes in the proximal region of the leaf, but has little effect on the distal region. These data support the heterochronic model of pea leaf morphogenesis proposed by Luet al. (International Journal of Plant Science157: 311–355, 1996).
... Natural and/or induced alleles are known in six genes that determine leaf morphology. Both loss of function (uni) and hypomorphic (uni-tac; lesser and TENDRIL-LESS (TL) genes (de Vilmorin and Bateson 1911;White 1917;Eriksson 1929;Lamprecht 1933Lamprecht , 1959Kujala 1953;Goldenberg 1965;Sharma 1972;Hofer et al. 2001;Taylor et al. 2001;Smirnova 2002;Kumar et al. 2004;Tattersall et al. 2005;Hofer et al. 2009). UNI, CRI, STP and TL are known to be transcription factors or coregulators of transcription factors; UNI, CRI and STP are respectively orthologues of LEAFY/FLORICAULA, ASYMMETRIC LEAVES 1/PHANTASTICA and UNUSUAL FLORAL ORGANS/FIMBRIATA (Hofer et al. 1997;Taylor et al. 2001;Tattersall et al. 2005;Hofer et al. 2009). ...
In angiosperms, leaf and stipule architectures are inherited species-specific traits. Variation in leaf and stipule sizes, and forms result from the interaction between abiotic and biotic stimuli, and gene regulatory network(s) that underlie the leaf and stipule developmental programme(s). Here, correspondence between variation in leaf and stipule architectures described for extant angiosperms and that induced mutationally and by imposition of stress in model angiosperm species, especially in Pisum sativum, was detected. Following inferences were drawn from the observations. (i) Several leaf forms in P. sativum have origin in fusion of stipule and leaf primordia. Perfoliate (and amplexicaul and connate) simple sessile leaves and sessile adnate leaves are the result of such primordial fusions. Reversal of changes in the gene regulatory network responsible for fusion products are thought to restore original stipule and leaf conditions. (ii) Compound leaf formation in several different model plants, is a result of promotion of pathways for such condition by gene regulatory networks directed by KNOx1 and LEAFY transcription factors or intercalation of the gene networks directed by them. (iii) Gene regulatory network for compound leaves in P. sativum when mutated generates highly complex compound leaves on one hand and simple leaves on other hand. These altered conditions are mutationally reversible. (vi) Simple leaves in model plants such as Arabidopsis thaliana despite overexpression of KNOx1 orthologues do not become compound. (v) All forms of leaves, including simple leaf, probably have origins in a gene regulatory network of the kind present in P. sativum.
... Mutants of unifoliata leaf morphology are also known in the leguminous plants Phaseolus vulgaris (Myers and Basett 1993), Vigna unguiculata (Fawole 2001) and Medicago truncatula (Wang et al. 2008). Besides UNI (UNI-TAC) and STP, several other genes such as AFILA (AF), INSECATUS (INS), MULTIFOLIATE- PINNA (MFP) and TENDRIL–LESS (TL) (de Vilmorin and Bateson 1911; Lamprecht 1959; Goldenberg 1965; Hofer et al. 2001; Smirnova 2002; Kumar et al. 2004) control meristematic activity in the rachis primordium and subprimordia for pinnae in the compound leaf differentiation in P. sativum. UNI gene mediates meristematic activity that grows rachis in proximodistal and mediolateral directions (Hofer et al. 1997; DeMason and Schmidt 2001; Gourlay et al. 2000; Yaxley et al. 2001; Prajapati and Kumar 2002; DeMason 2005; Mishra et al. 2009; Kumar et al. 2010); these activities are respectively made determinate by AF and INS and TL and MFP (DeMason and Schmidt 2001; DeMason and Chawla 2004a and b; Mishra et al. 2009; Kumar et al. 2010). ...
UNIFOLIATA [(UNI) or UNIFOLIATA-TENDRILLED ACACIA (UNI-TAC)] expression is known to be negatively regulated by COCHLEATA (COCH) in the differentiating stipules and flowers of Pisum sativum. In this study, additional roles of UNI and COCH in P. sativum were investigated. Comparative phenotyping revealed pleiotropic differences between COCH (UNI-TAC and uni-tac) and coch (UNI-TAC and uni-tac) genotypes of common genetic background. Secondary inflorescences were bracteole-less and bracteolated in COCH and coch genotypes, respectively. In comparison to the leaves and corresponding sub-organs and tissues produced on COCH plants, coch plants produced leaves of 1.5-fold higher biomass, 1.5-fold broader petioles and leaflets that were 1.8-fold larger in span and 1.2-fold dorso-ventrally thicker. coch leaflets possessed epidermal cells 1.3-fold larger in number and size, 1.4-fold larger spongy parenchyma cells and primary vascular bundles with 1.2-fold larger diameter. The transcript levels of UNI were at least 2-fold higher in coch leaves and secondary inflorescences than the corresponding COCH organs. It was concluded that COCH negatively regulated UNI in the differentiating leaves and secondary inflorescences and thereby controlled their sizes and/or structures. It was also surmised that COCH and UNI (LFY homolog) occur together widely in stipulate flowering plants.
Induced mutagenesis is now an established method for crop improvement. Mutation techniques by using ionizing radiations and other mutagens have successfully produced quite a large number of new promising varieties in different plant species. Since beginning there are step by step improvement in technical procedure for application of induced mutation for crop improvement and voluminous knowledge have developed for successful and accurate application of the technique. The chapter highlights a bird’s eye view of the prospects, procedures, possibilities and problems of mutation breeding. There are huge publications by many workers highlighting their success and failure on induced mutagenesis. Attempt has been made to highlight different important basic aspects which may be helpful as guideline for large scale mutagenesis work on any ornamental crop. Author tried to put together all available information on mutagens and dose to develop a complete documentation of the results of the research conducted by different scientists over the last about 80 years.
Present scenario of global warming has impact on the overall progress of mankind from agriculture sector to health sector. Abiotic stress due to climate change has declined crop yield and production worldwide. This decline in crop production has impact on agriculture sector which finally affected the human being by scarcity of food supply. According to the survey conducted, by the end of 2050 global population is expected to cross 9.7 billion. The increased population will create pressure on the present limited natural resources to produce more food. For feeding the increasing population there is need to produce almost 50 per cent more food, feed, and bio-fuel. Breeders across the world are indulged in developing smart crops by utilizing both conventional and modern breeding techniques. The most important breeding approach is identification of novel quantitative trait loci/candidate gene which can be used further for improvement of breeding program of a particular crop. Pisum is one of the most important vegetable crops consumed globally. Like in other crops, QTL mapping, gene mapping, and association mapping for various traits have been studied in Pisum crop. Here, in this chapter, we will discuss the importance of different mapping in vegetable crops and provide a detail of identified QTLs/candidate genes for various traits in Pisum.
The timing of developmental phase transitions is crucial for plant reproductive success, and two microRNAs (miRNA), miR156 and miR172, are implicated in the control of these changes, together with their respective SQUAMOSA promoter binding–like (SPL) and APETALA2-like (AP2) targets. While their patterns of regulation have been studied in a growing range of species, to date they have not been examined in pea (Pisum sativum), an important legume crop and model species. We analysed the recently released pea genome and defined nine miR156, 21 SPL, four miR172, and five AP2-like genes. Phylogenetic analysis of the SPL genes in pea, Medicago and Arabidopsis confirmed the eight previously defined clades, and identified a ninth potentially legume-specific SPL clade in pea and Medicago. Among the PsSPL, 14 contain a miR156 binding site and all five AP2-like transcription factors in pea include a miR172 binding site. Phylogenetic relationships, expression levels and temporal expression changes identified PsSPL2a/3a/3c/6b/9a/9b/13b/21, PsmiR156d/j and PsmiR172a/d as the most likely of these genes to participate in phase change in pea. Comparisons with leaf morphology suggests that vegetative phase change is unlikely to be definitively marked by a change in leaflet number. In addition, the timing of FT gene induction suggests that the shift from the juvenile to the adult vegetative phase may occur within fourteen days in plants grown under inductive conditions, and calls into question the contribution of miR172/AP2 to the floral transition. This work provides the first insight into the nature of vegetative phase change in pea, and an important foundation for future functional studies.
Pea leaf determination was examined by culturing excised leaf, leaflet, and tendril primordia of different ages on a nutrient medium. Pinna primordia were designated as 1) determined, if they grew normally in culture; 2) undetermined, if they grew into differentiated structures that were morphologically and anatomically different from either leaflet or tendril; or 3) partially determined, if the two pinnae of an opposite pair developed unequally in isolation, or for leaflet pinnae only, if laminae were initiated but did not develop completely. The compound pea leaf as a whole is determined over four plastochrons of development. Proximal pinnae are determined during the second leaf plastochron, approximately 0.8 plastochron after their initiation. The second most proximal pair of pinnae is determined during the third plastochron, and the terminal portion of the rachis is determined last, during the fourth plastochron. Determination of leaflet dorsiventrality is gradual, requiring a critical minimum period with the leaf in physiological contact with the shoot system. The rachis primordium, when isolated from the shoot, does not affect determination of its pinnae as leaflets or tendrils. Afila and tendril-less homeotic mutations do not alter the timing of pinna determination.
After gamma-ray treatment of pea seeds, a mutation tendril-less2 (tl2) was obtained. In heterozygote, it transforms tendrils into a narrow leaflets, resembling an action of the classic mutation tl. In contrast to tl, the novel mutation, when homozygous, does not affect leaf development but supresses pigmentation of petals. It is suggested that both genes (Tl and TI2), responsible for transformation of leaves into an organ of climbing, have common origin from a gene controlling anthocyanine synthesis
Experimental evidence has shown that all forms of peas previously described as species have a diploid chromosome number of 14, that no sterility barriers exist, and that gene exchange is complete. The genus Pisum is therefore best regarded as monospecific in accordance with Lamprecht’s (1966) view. He classified the different forms as ecotypes included under Pisum arvense Linné, the wild-growing form of the two described by Linné. The ecotypes abyssinicum Braun; arvense (Linné) Lamprecht (including elatius Steven, jomardi Schrank, and transcaucasicum Stankov); fulvum Sibthorp and Smith; and humile Boissier (including syriacum/Berger/Lehmann) occur as wild-growing populations. All man-made genetic variations were collected together under the name sativum, the domesticated race. This system of classification is practical and workable, though perhaps not taxonomically orthodox. Pisum formosum Steven, which is a tuber-forming perennial, was separated to form the genus Alophotropsis (Boissier) Lamprecht.
To reveal the role(s) of stipules, pea genotypes of nearly common genetic background, but differing in the combination of wild type and mutant alleles of STIPULE-REDUCED, AFILA and TENDRIL-LESS genes together with COCHLEATA gene, were grown in a field in completely randomized design during the winter-spring seasons through 2006 to 2011 at the farm of the institute. The genotypes were characterized organ-wise, for morphological and anatomical features, photosynthesis and biomass accumulation and partitioning. Stipules were found similar to leaves/leaflets in respect of stomata shape, size and frequency as well as size and shape of pavement cells in epidermis and internally, in possession of palisade- and spongy-mesophyll parenchyma and veins in high density. In contrast to leaflets, which possess single primary vein and smooth lamina margin, the stem proximal part of stipule had serrations and possessed several primary veins; however, its distal part had single primary vein and smooth lamina margin. Simultaneous occurrence of the properties of simple and pinnately compound and palmate lamina structure were identified in stipules. Compared to leaflets in which the highest order of veinal reticulation was quinternary that in stipule was sexternary. Stipules were highly effective photosynthetic organ, responsible for about 30% of photosynthesis in phytomeres. Stipules enlarged plant's photosynthate/ biomass production and accumulation capacity by about 72%. The presence of stipules avoided feedback inhibition of photoassimilation in leaves, by ensuring that pods/seeds remained the principal sink for photoassimilates. The presence of stipules was essential for high harvest index and grain yield; the harvest index in the stipule-deficient genotypes was 2.8 fold lower as compared to genotypes possessing wild type stipules. The plants in which the stipules were intact but leaves had been surgically removed, produced seeds proving that stipules alone were sufficient for survival and completion of life cycle in pea plant. The plants from which stipules had been surgically removed had reduced growth indicating that the presence of stipules was vital for continued shoot growth. The observations altogether demonstrated additive interaction between stipules and leaves in the accomplishment of photosynthesis, accumulation of biomass and the production of seeds in high yield with high harvest index. It was concluded that the presence of photosynthetically efficient stipules is a requirement in the breeding of high yielding cultivare of the pea (Pisum sativum) crop.
In the present study, two pea (Pisum sativum L.) mutants experimentally obtained after gamma - irradiation of dry seeds were described. Both mutants were characterized with enlarged leaf size. The mutant 2/462 was shown to have semi - dominant inheritance pattern; 2/927 was sterile and inherited monogenic recessive. The induced mutations have pleiotropic effect affecting morphological and reproductive traits. New mutants had similar phenotypes to previously named mutants latifolium (lat) and cabbage leaf (calf), but no allelism tests were made between the new and the previously reported mutants. Both mutant lines may be useful plant material for research on leaf development.
In grain pea Pisum sativum, 16 genotypes constructed by combining wild type and mutant alleles of MULTIFOLIATE PINNA (MFP),AFILA (AF), TENDRIL -LESS(TL) and UNIFOLIATA (UNI) genes, which differed in leafblade morphology, were quantitatively phenotyped for allometry. The biomass partitioning among root, stem, stipule, leaf blade and seeds was unique for these genotypes suggesting that MFP AF TL and UNI genes determined leaf blade and plant architecture. Gene actions were inferred on the basis of mutant phenotypes. Biomass of all the organs was found to be increased in the tl single mutant. The afmutation singly and in combination with mfp, mfp tl, mfp uni-tac or mfp tl uni-tac decreased biomass of all the organs. Allocation of biomass to leaves was increased at the expense of that to seeds or seeds and stems by a single mfp mutation or in combination with uni-tac, aftl, tl uni-tac and aftl uni-tac mutations. The AF and MFP functions are essential in pea cultivars for high yield of grains.The mechanism for simultaneous control of leaf blade and plant architecture suggested by mutant phenotypes has three elements. The MFP, AF TL and UNI genes exercise control over meristematic activity in all the organs. Their determination of leaf blade morphology and size affect net photosynthesis or metabolite supply. The quantities of available metabolites determine numbers and sizes of organs or partitioned total biomass. The tl allele is identified as a genetic marker/determinant for breeding tendril-less prolific pea cultivars for obtaining herbage and grains in high yields.
The way of inheritance type of the pea leaves is a parameter of study in this investigation. Bulgarian and foreign varieties and their hybrids with different quality and quantity traits are used. Biological and agronomic characteristics are worked out. In genotype's crossover between acacia and afila type of the leaves is observed double recessive inheritance on pleiofila and mon-ogenetic recessive on afila type. Do not established inter-allel interactions of the genes determined type of the leaves in investigated forms. A large diversity for different traits is received in hybrid populations. The pleiofila hybrids represent selection interest due to their high productivity of pod and seeds per plant that is prerequisite for future selection work.
Study Czech collection of pea showed very low representation of germplasm from Armenia, Georgia, Mongolia, Tajikistan, Morocco and Algeria, the areas designated as centres of genetic diversity of pea with the occurrence of economic characteristics that are of current cultivars due to intensive breeding completely disappeared. 68 pea accessions were obtained from gene bank VIR St Petersburg. As a control cultivars were chosen cultivars of field peas (Pisum sativum L. ssp sativum var. sativum) - Sponsor and Zekon and fodder pea (P. sat. L. ssp arvense) - Arvika.
Statistically significant positive correlation (r = 0.78) between leaflet margin shape in the second real leaves and leaflet margin shape in the first flowering node, character of antocyanin spot and wings (r = 0,93) and vexillium colour (r = 0,91) was confirmed.
The Euclidean distance formula for calculate the coefficient of similarity was elected.
The minimum value (1.41) was found between Kamyshlovskyj mramornyj myelkyj 1584 and Untitled 999 accessions. The maximum value of the coefficient of similarity (17.07) was found between Primavil and K 38_2314 accessions. High coefficient of similarity was found between the control cultivar Sponsor and 12 accessions. In control cultivar Arvika differing from cultivar Sponsor by flower colour and type of growth was found 12 accessions with high coefficient phenotypic similarity.
With our developed method for the molecular analysis of eIF4E resistance gene PSbMV P1 virus were performed PCR analysis of length polymorphism of intron 3 using primers 750F + 586 g-R. Any accessions containing a known resistance allele to PSbMV were detected. At the sample no. 1589 was amplified PCR product suggesting the presence of a novel allele. This was confirmed by sequencing analysis of the product. Direct virological testing confirmed that this is new allele resistance PSbMV.
Recent work on the venation patterning and morphogenesis of leaf/leaflet has posed the question how different are these in tendrils, which are another type of vegetative lateral organ. Here, the venation patterns of leaflets, stipules and tendrils were compared in the model species, P. sativum. Unlike reticulated venation in leaflets and stipules, venation in tendrils comprised of one or more primary veins. A few secondaries were attached to a primary vein, mostly distally. Bilaterally symmetrical secondary veins were rare. The primary veins in tendrils were daughter strands from dichotomously divided mother veins in rachis, connected finally to vascular strands in stem. A tendril received primary vein from one or more mother strands. Some mother strands contributed primary veins to proximal, distal and terminal domain tendrils of af leaf. The tendrils shared the multi-primary vein character with stipules. Vein redundancy provided a mechanism for survival of tendril/leaf against injury to some of the veins/mother veins. The presence of aborted primary veins that did not reach apex, rows of cambium cells attached to primary vein(s) at apex, the pattern of attachment of primary veins to mother veins and cessation of vein growth in apical direction in aborted tendrils of af lld genotype indicated that the growth of primary veins and tendril was acropetal. Loss-of-function of AF extended the repression of TL and MFP genes on leaflet development from distal and apical domains to proximal domain of leaves in af mutants.
In the wild type P. sativum, each of the adult plant stem nodes, bears a pair of sessile foliaceous stipules and a petiolated unipinnately compound leaf of 4 to 6 leaflets and 7-9 tendrils. The stipule-reduced (st) and cochleata (coch) single null mutants and coch st double null mutant differ fom the wild type in respectively having sessile stipules of much reduced size, petiolated simple and/or compound leaf-like stipules and no stipules. It is also known that coch leaves are somewhat bigger than st and wild type leaves. Here, pleiotropic phenotype of coch st double mutant was investigated. The morphologies of stipules and leaf were quantified in the field grown plants and microcultured shoots, latter in the presence and absence of gibberellic acid and N-1-naphthylphthalamic acid. The observations showed that as compared to the corresponding plants or shoots of COCH ST (WT) genotype, (a) coch st plants bore leaves in which all the organs were hypertrophied; (b) full complement of leaflets and 3-5 tendrils were formed on leaf; (c) the microcultured coch st shoots were taller despite lower number of nodes, and (d) they also produced leaves in which all the organs were bigger and the ratio of leaflets/tendrils was higher. It was concluded that in coch st double mutant (a) ST function is essential for stipule primordium differentiation, in the absence of COCH function and (b) absence of negative feedback loops between simple stipules and compound leaf for metabolite utilization allows hypertrophied growth in leaves.
Gene gf (for greenish flower colour) and gene efr (for early flowering) of Pisum sativum were combined with other mutant genes of the genome. Their behaviour in different combinations was studied with regard to the seed production of the various recombinants. The following results were obtained:1. Gene gf of mutant 189 was tested in 22 combinations. Some recombinant types, homozygous for gf, for genes for long or very long internodes and for stem fasciation, show a seed production of more than 800% of that of mutant 189.2. Gene efr was tested in 59 combinations. The seed production of a high proportion of the recombinants studied was found to be essentially better than that of the parental genotype. This holds true for both long- and short-stemmed recombinants.3. An early-flowering tendril-less super-dwarf shows high productivity partly due to the positive alteration of the pleiotropic pattern of gene efr.4. The selection value of both the genes could be strongly improved by combining them with specific other mutant genes of the Pisum genome.
The leaf blade of Pisum sativum L. possesses proximal leaflets and distal tendrils and is altered by two recessive mutations that affect pinna identity, afila (af) and acacia/tendrilless (tl). Using morphological and histology features we characterized the variation in leaf form and pinna anatomy of the Af and Tl heterozygous genotypes. Our goal was to identify the specific interactions of these genes and the relative influence of each in regulating all structural components of the leaf and pinna phenotypes. The Tl/tl genotypes possess broad tendril - narrow leaflets in the distal region that are histologically more similar to leaflets than to tendrils. The afafTltl leaves have simple tendrils in the distal region and branching complexity that is intermediate between that of aftl and af leaves in the proximal region. Only the most distal tips are slightly expanded. Because the Af and Tl genes interact to affect almost all aspects of leaf and pinna form, they affect development at multiple levels of organization.Key words: acacia, afila, Fabaceae, leaf morphology, leaf anatomy, Pisum sativum.
The leguminous flora of Delhi comprises 78 Papilionoideae, 24 Caesalpinioideae and 24 Mimosoideae species; 80 of them are perennials. Five types of imparipinnate and two types of paripinnate compound leaves was observed on the species. The paripinnate leaves are bipinnate in 25 species (mostly mimosoid) and bifoliate in two species. The imparipinnate leaves were trifoliate or multifoliate in 59 papilionoid species and multifoliate in 16 caesalpinioid species; four of the papilionoid species produced leafletted and tendrilled unipinnate leaves. Leaves were bifacially simple in 22 species, simple with ectopic terminal growth in one species and simple tendril in one species. Twenty-one species (mostly mimosoid) were devoid of stipules. In 82 species stipules were small and free. Stipules were large and lobed in 17 species and large and adnate in four species. Two species of Caesalpinioideae produce compound leaf-like stipules. All four stipule phenotypes of 126 species corresponded with stipular phenotypes observed in wild type, coch, st and coch st genotypes of the model legume P. sativum. The seven leaf phenotypes observed in 126 species corresponded with phenotypes expected among combinations of uni (uni-tac), af, ins, mfp and tl mutants of P. sativum and sgl1, cfl1, slm1 and palm1 mutants of M. truncatula, also an IRL model legume. All the variation in leaf and stipule morphologies observed in the leguminous flora of Delhi could be explained in terms of the gene regulatory networks already revealed in P. sativum and M. truncatula. It is hypothesized that the ancestral gene regulatory networks for leaves and stipules produced in Leguminosae were like that prevalent in P. sativum.
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Premise of the study:
Processes of leaf morphogenesis provide the basis for the great diversity of leaf form among higher plants. The common garden pea (Pisum sativum) offers a developmental model system for understanding how gene and hormone interactions impart a large array of mutant leaf phenotypes. •
Methods:
To understand the role of auxin in AF and UNI gene function and their interaction, we compared the range of leaf phenotypes on afila (af) and unifoliata (uni) double mutants, examined the effects of these mutations on auxin levels, auxin transport, auxin response via DR5::GUS, and expression of auxin-regulated genes. •
Key results:
The adult leaves of af uni double mutants have leaflets and tendrils and typically possess two lateral pinna pairs and a terminal leaflet. The af mutants have higher auxin content, stronger auxin response, and higher expression of auxin responsive genes than wildtype. The uni mutant has reduced auxin content and transport, whereas the uni-tac mutant has higher auxin content and transport and reduced auxin response compared to wildtype. •
Conclusions:
Auxin concentration and response differences characterize the antagonistic relationship between AF and UNI in pea leaf development. The mechanism involves modulation of auxin mediated by one or both genes; UNI is expressed in and promotes high auxin levels, and AF suppresses auxin levels.
Pea has been grown for food and feed since its domestication the Mediterranean region in approx. 7000 B.C. It serves as an excellent source of protein, fiber, minerals and vitamins especially in developing countries. Nitrogen fixation by Rhizobium leguminosarum in symbiotic relationship with pea is a primary benefit to inclusion in crop rotations. Several biotic and abiotic stresses constrain production worldwide. Genetic resistance is available for many of the diseases and when coupled with cultural control methods can be effectively controlled. Heterosis and male sterility are present in pea, however, to date no hybrid cultivars have been developed or commercialized due to difficulty associated with cross pollination. Seed production and maintenance of inbred pure lines is through bulk production or single plant selection in disease free fields isolated by at least 30 m from neighboring pea fields. Clean equipment, routine field inspection and use of electric eye technology further increase seed purity. Wide adaptation and use of modern technology in cultivar improvement ensure that pea will continue to be an important crop worldwide.
The yield potential, quality and level of resistance to powdery mildew (Erysiphe pisi DC.) of afila smooth seeded pea (Pisum sativum L.) were tested in the field trials. The cultivars and breeding lines Mozart, Consort-R, AGT-01, Cebe-co 1171 and AGT-GH surpassed the control cv. Gotik in the yields of dry seed, in contrast the dry seed yields of Highlight, AGT-KR, Melfort and LU 390-R2 were about 12–27% lower than that of the control. The low seed yield was caused by virus infections (PEMV, BYMV), root diseases (Pythium, Fusarium), and a low level of thousand seeds weight (TSW). Material crossing with donors possessing high yield potential, a higher TSW, and a higher tolerance to root dis-eases had a positive effect on the dry seed yield. The main objective of the resistant pea breeding programme is afila smooth seeded pea resistant to powdery mildew, with a high tolerance to viruses, root diseases, and lodging, with the stem length of 60 to 75 cm, and with high yield potential.
Shoot-like compound leaves are ancient and predate the evolutionary origin of whorled flowers. By studying leaves of this type it may be possible to gain further insight into similar fundamental processes in plant development. Here, we describe the pea compound leaf from the viewpoint that considers it as a determinate lateral shoot. By integrating research on developmental mutants, we present a model of the genetic control of patterning in the pea leaf.
The pea Pisum sativum genes AFILA (AF), TENDRIL-LESS (TL) and UNIFOLIATA (UNI) have been shown previously to be involved in the compound leaf blade patterning and AF and TL genes in leaf blade growth. Here, the interactions of the previously described UNI gene with AF and TL were studied using a new uni-tac allele and af and tl alleles, in the new background of cultivar Pusa Harbhajan. The leaf blades formed on the embryonic and mature pre- and post-flowering leaves from plants of AFTLUNI, afTLUNI, AFtlUNI, AFTLuni-tac, aftlUNI, afTLuni-tac, AFtluni-tac and aftluni-tac genotypes were measured for pinna growth in terms of their laminated and non-laminated organs. The investigation showed that UNI activates, and AF and TL repress the leaf blade ramification. UNI and AF act together in lamina-dominated pinna growth; we call this the AF-dependent leaflet development pathway. Presence of second AF-independent pathway for leaflet formation is described. TL and UNI are identified as antagonists of such pathway(s). UNI, AF and TL functions can up-regulate as well as down-regulate leaf-blade morphogenetic developmental events, in organ-specific manner.
The pleiofila phenotype (afaftltl double mutant) of Pisum sativum arises from two single-gene, recessive mutations known to affect the identity of leaf pinnae, afila (af), and acacia (tl). The wild-type leaf consists of proximal leaflets and distal tendrils, whereas the pleiofila leaf consists of branched pinnae terminating in small leaflets. Using morphological measurements, histology, and SEM, we characterized the variation in leaf form along the plant axis, in leaflet anatomy, and in leaf development in embryonic, early postembryonic, and late postembryonic leaves of aftl and wild-type plants. Leaves on aftl plants increase in complexity more rapidly during shoot ontogeny than those on wild-type plants. Leaflets of aftl plants have identical histology to wild-type leaflets although they have smaller and fewer cells. Pinna initiation is acropetal in early postembryonic leaves of aftl plants and in all leaves of wild-type plants, whereas in late postembryonic leaves of aftl plants pinna initiation is bidirectional. Most phenotypic differences between these genotypes can be attributed to differential timing (heterochrony) of major developmental events.
We examined a series of eight pea genotypes differing in three naturally occurring allelic mutations, i.e., af (afila), st (stipules reduced), and tl (tendril-less) and three species, five cultivars, and one interspecific hybrid of tomato differing in SP (SELF-PRUNING) allele composition to determine whether different phenotypes ontogenetically express different biomass partitioning patterns compared to the isometric partitioning pattern and an interspecific 3/4 scaling "rule" governing annual growth with respect to body mass. The slopes and "elevations" (i.e., α and log β, respectively) of log-log linear regression curves of bivariate plots of leaf, stem, and root dry mass and of annual growth vs. total body mass were used to assess pattern homogeneity. The annual growth of all pea and tomato phenotypes complied with the 3/4 growth rule. The biomass partitioning patterns of all tomato phenotypes were statistically indistinguishable from the isometric pattern as were those of the pea wild type and three single-mutant genotypes. However, significant departures from the isometric (and pea wild type) biomass allocation pattern were observed for three genotypes, all of which were homozygous for the af allele. These results open the door to explore the heritability and genetics underlying the allometry of biomass partitioning patterns and growth.
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