The Effects of Shading on the Yield Components of Six 'Leafless' Pea Genotypes

The effect of unshaded, 30 and 80 per cent shade conditionson the growth rate and yield composition of six ‘leafless’pea (afaf: stst) genotypes has been determined. No evidencefor a photoassimilate source limitation was found in the unshadedand 30 per cent shade treatments, but 80 per cent shade significantlyreduced the yield and the rate of d. wt and node productionof all the genotypes. Yield development was followed in each environment by determiningthe number of flowering nodes, the number of flowers producedper node, the pod number per node and the seed number per pod.The theoretical maximum for each of these yield components wasdetermined and compared with the actual level realized by theplant. Approximately 50 per cent of the theoretical potentialof each yield component was realized and this appeared to beindependent of the shade treatment. This stepwise reductionin potential yield throughout the development of the plant wassimilar to that reported for conventional pea plants. Pisum sativumL, ‘leafless’ peas, shading, growth rate, yield'components, source limitation     

Residual effect of germination temperature on the growth of peas

Summary In Pisum sativum there is a marked difference in the final height of the plants depending on the temperature at which the seed is germinated. Significant differences were also found in such parameters as growth rate, node of first flower, seeds per plant, pods per plant as well as flowers per plant.     

Flowering in Pisum: Effect of the Parental Environment

The parental photoperiod appears to have no effect on the floweringnode of the progeny, provided the progeny seed is selected tobe of the same weight. However, the parental photoperiod doesinfluence the mean seed weight and seed weight is shown to causesignificant alterations in the flowering node and time as wellas causing alterations in vegetative characters like the internodelength and the rate of leaf expansion and node formation. Theeffect of seed weight on flowering is thought to result fromthe alterations in the growth rate. On the other hand, vernalization of the parents does appearto cause a small, but significant effect on the flowering nodeof the progeny (i.e. it could be transmitted through a meioticdivision). This effect disappears in the next generation andthe possible nature of this effect is discussed. Pisum sativum L, garden pea, flowering, photoperiod, vernalization     

Flowering in Pisum: the Effect of Age on the Gene Sn and the Site of Action of Gene Hr

Late cultivars of peas behave as quantitative long day plants.The reason that they flower between nodes 20 and 35 under an8 h photoperiod is shown to be because the leaves and maturestem produce a more promotory ratio of the flowering hormonesas they age. Later formed leaves may also start with a slightlymore promotory ratio than the leaves produced at a lower node.The gene Sn controls the production of a flower inhibitor andit is suggested that the activity of this gene in a leaf isgradually reduced as the leaf ages. From grafting experiments,the site of action of the gene Hr is shown to be in the leavesor mature stem and not at the shoot apex. This supports a previoussuggestion that the gene Hr is a specific inhibitor of the ageingresponse of gene Sn. Gene Hr is shown to cause a substantial delay in the floweringnode of decotyledonized plants of genotype If e sn hr undershort day conditions, suggesting that Hr has little effect inthe cotyledons. It is argued that the gene sn is a leaky mutantand that gene Hr does not control a photoperiod response inits own right but has its effect through the Sn locus. From a comparison of intact plants and self-grafts of the lategenotype If e Sn hr it is shown that under the conditions usedphysiological age may be of more importance than chronologicalage in determining flowering in peas. Reasons for the smalleffect of defoliation treatments on flowering are discussedas well as possible reasons for the promotory effect of decotyledonizationon the flowering node of late lines. Pisum sativum L, flowering, ageing, genetic control     

Seed Production in Leafless and Conventional Phenotypes of Pisum sativum L. in Relation to Water Availability Within a Controlled Environment

Water requirements in relation to seed production was studiedin near-isogenic lines of leafless (afafstst) and conventional(++++) pea plants (Pisum sativum). The plants were grown toseed maturity in pots in a controlled environment under conditionsof high, medium and low irrigation levels. When each genotypewas irrigated independently and on demand and the soil moisturecontent maintained at 65–80 per cent of full capacitythere was no significant phenotypic difference in water useefficiency (WUE), defined as g d. wt seed per kg H₂O utilized.There existed genotypically-controlled upper and lower limitsto yield between which the total dry weight of seed per plantcan be determined by water availability. There was no significantdifferential effect of genotype or of irrigation treatment onthe number of pods, number of seed, unit seed dry weight andbiological yield per plant. There was significant interactionon stem length, and leaf area at specified nodes. When the wateractually required in relation to the water available was takeninto account, the leafless phenotype consistently utilized 33–38per cent less water and produced a correspondingly lower totaldry weight of seed than the conventional counterpart. Independentlyof regime the total dry weight seed per phenotype remained anear constant proportion of the above-ground biomass. Pisum sativum L., garden pea, leafless peas, seed production, water availability     

Time of Flowering of Pea (Pisum sativum L.) as a Function of Leaf Appearance Rate and Node of First Flower

In order to assess the influence of environmental conditionson time of flowering of pea (Pisum sativum L.), a serial sowingtrial was conducted over 2 years at Dijon, France, on two wintercultivars Frisson and Frilene. Time of flowering was analysedaccording to two variables: the leaf appearance rate R_L andthe node of first flower N_I. R_L was linearly related to temperature (r² = 0·94). Thebase temperature was 2°C for both varieties. Growth rateaccounted for the residual variability of R_L . Photoperiod andtemperature acted on N_I in an additive way. Frilene, the latergenotype, was more responsive than Frisson. A model for predicting time of flowering based upon these resultsis proposed. Deviations from this model were related to N nutritionin interaction with the plant water relations. Steps for improvingthe model are then discussed.Copyright 1993, 1999 Academic Press Pisum sativum L., pea, flowering, temperature, photoperiod, phyllochron, model     

Apical Senescence in Pisum: A Direct or Indirect Role for the Flowering Genes ?

Apical senescence was examined in a range of intact and defloweredflowering genotypes under both long and short photoperiods.The flower inhibitor produced by the gene Sn, appears to havea direct effect on apical senescence since it can delay apicalsenescence under short day conditions in the absence of flowerand fruit development or where the rate of such developmentis the same in different treatments. Gene Hr can magnify thiseffect. Gene E, on the other hand, appears to influence apicalsenescence only indirectly through the effect it has on flowerand fruit development. The flowering genes at the If, sn andhr loci are also thought to have indirect effects on apicalsenescence. Even in deflowered plants apical senescence appearsto occur eventually in continuous light in all genotypes testedindicating that the presence of developing fruits, althoughpromotory, is not essential for apical senescence. Pisum sativum L., garden pea, flowering, senescence     

Cost of reproduction in a spring ephemeral species, Adonis ramosa (Ranunculaceae): carbon budget for seed production

BACKGROUND AND AIMS: Spring ephemerals have a specific life-history trait, i.e. shoot growth and sexual reproduction occur simultaneously during a short period from snowmelt to canopy closure in deciduous forests. The aim of this study is to clarify how spring ephemerals invest resources for seed production within a restricted period. METHODS: In order to evaluate the cost of reproduction of a typical spring ephemeral species, Adonis ramosa, an experiment was conducted comprising defoliation treatments (intact, one-third and two-thirds leaf-cutting) and fruit manipulations (control, shading and removal) over two growing seasons. In addition, measurements were made of the movements of carbon assimilated via (13)C tracing. KEY RESULTS: Survival rate was high irrespective of treatments and manipulations. The proportion of flowering plants and plant size decreased as a result of the defoliation treatments over 2 years, but the fruit manipulations did not affect flowering activity or plant size. Seed set and seed number decreased as a result of fruit shading treatment, but the defoliation treatments did not affect current seed production. Individual seed weight also decreased in the second year due to fruit shading. The (13)C tracing experiment revealed that young fruits had photosynthetic ability and current photosynthetic products from leaves were mainly transferred to the below-ground parts, while translocation to fruit was very small even when fruit photosynthesis was restricted by the shading treatment. CONCLUSIONS: Current foliage photosynthetic products are largely stored in the below-ground parts for survival and future growth, and about one-third of the resources for seed production may be attained by fruit photosynthesis. Therefore, the trade-off between current seed production and subsequent growth is weak. The cost of seed production may be buffered by sufficient storage in the below-ground organs, effective photosynthesis under high irradiation and self-assimilation ability of fruits.     

Factors affecting fruit and seed production of Paeonia ostii “Feng Dan”, an economically important oil tree

Many factors may affect reproduction of animal-pollinated species. In this study, the effects of pollen limitation, attractive traits (flower number, plant height and flower width) and flowering phenological traits (flowering onset, duration and synchrony) on female reproduction, as well as the patterns of variation in fruit and seed production within plants, were investigated in Paeonia ostii “Feng Dan” over two flowering seasons (2018 and 2019). Fruit set was very high (90%), and pollen supplementation did not increase fruit and seed production in either year, indicating no pollen limitation. Fruit set, ovule number per fruit and mean individual seed weight per fruit were not affected by any of the six attractive and phenological traits in either year, whereas seed number per fruit was related to the three attractive traits in one or both years. Seed number per plant was positively affected by the three attractive traits and best explained by flower number in both years, but the effect of each of the three phenological traits on seed number per plant differed between years. Within plants, the fruit set, ovule number, seed set and seed number per fruit declined from early- to late-opening flowers, presumably because of resource preemption, but the mean individual seed weight did not vary across the flowering sequence. Our study shows that attractive traits of Paeonia ostii “Feng Dan” are more important than flowering phenological traits in the prediction of total seed production per plant.     

大花百子莲的结实和结籽格局及种子产量影响因素分析

在2007和2008年对大花百子莲(Agapanthus praecox subsp.orientalis‘Big Blue’)在群体、果序和单果水平上的结实和结籽格局进行了观测,并依据每花序单花数、单花胚珠数和单粒种子质量推算其单株潜在种子产量;此外,采用人工补充授粉方法研究了不同授粉方式对大花百子莲结实和结籽的影响,并对其种子产量的影响因素进行了分析。结果表明:从始花期、盛花期至末花期,随开花时间的延迟,大花百子莲植株的每花序单花数和座果率均逐渐降低,其中始花期开花的植株每花序单花数和座果率均最高。从果序基部、中部至顶端,座果率和结籽率均逐渐减小且差异显著。在果序基部和中部,果实内种子败育率沿果实基部至顶端依次降低,而单粒种子质量则依次增加;且果序基部的果实各部位的种子败育率均低于果序中部的果实,而单粒种子质量则差异不大。大花百子莲的单株潜在种子产量约为43.18 g,而其单株实际种子产量约为2.87 g,远低于其潜在种子产量。自然授粉的大花百子莲每柱头花粉数为27.5粒,而自花授粉、同株异花授粉和异株授粉的每柱头花粉数均超过100粒;与自然授粉、自花授粉和同株异花授粉植株相比,异株授粉植株的座果率和结籽率均显著增加。研究结果显示:大花百子莲的结实和结籽格局呈现非随机分布规律,在引种地存在有性繁殖障碍;养分和授粉状况是制约大花百子莲种子产量的重要因素。     

Metabolism of Phloem-borne Amino Acids in Maternal Tissues2Pisum sativum L.)

The amino acid composition of the EDTA-induced phloem exudatereaching the fruit and the seed, and of the solutes releasedby the seed coat during fruit development were determined inglasshouse-grown pea (Pisum sativum L. cv. Finale) suppliedeither with nitrate-free nutrients (nodulated plants) or withcomplete medium (non-nodulated plants). The EDTA-promoted exudationtechnique was used supposedly to collect phloem sap and theempty seed technique supposedly to collect the solutes secretedby the seed coat to the embryo sac cavity. In young seeds embryosac liquid was sampled directly from the embryo sac. The maincarbohydrate transported and secreted was sucrose. The mainamino acids reaching the fruit were asparagine, glutamine, andhomoserine. Their proportions were steady during a day-nightcycle and throughout fruit development. Amino acid compositionchanges occurred first in the pathway from fruit stalk to seedfunicle, due to the formation of threonine (probably from homoserine)and in the seed coat due to production of glutamine, alanineand valine which, together with threonine were the main secretedamino acids. The temporary nitrogen reserves of the pod walland seed coat were remobilized as asparagine during senescence.Phloem exudate of nodulated plants showed a higher (about twice)proportion of asparagine but lower proportions of homoserineand glutamine than in EDTA-induced phloem exudate of nitrate-fedplants. The two types of nitrogen nutrition also produced somechanges in relative proportions of threonine and homoserinesecreted by the seed coat. Key words: Pisum sativum, phloem, amino acids, pod wall, seed coat     

The Photosynthetic and Respiratory Potential of the Fruit in Relation to Seed Yield of Leafless and 'Semi-leafless Mutants of Pisum sativum L.

The photosynthetic and respiratory net CO₂ exchange potentialof the fruit of Pisum sativum was evaluated in one normal andthree mutant genotypes differing widely with respect to foliagearea and morphology: AfAf.StSt, conventional; AfAf.StSt, vestigialstipules; afaf.St.St, all leaflets replaced by tendrils butstipules normal (here termed ‘semi-leafless’) andafaf.stst, tendrils and vestigial stipules (‘leafless’).Agronomically the latter phenotype offered improvement in resistanceto lodging, crop drying, and harvester through-put. On a dry weight basis, fruits of the leafless mutant were consistentlymore active photosynthetically in terms of CO² uptake from theatmosphere during the initial 18 days post-anthesis than werethe corresponding fruits of the other three phenotypes. Duringthe subsequent 16-day period of seed filling there was no markeddifference between phenotypes and the fruit continuously lostCO² to the atmosphere, but significantly less CO² was lost inthe light (40 k lux) than in the dark. That the fruit of theleafless mutant may therefore benefit from the increase in lightavailable to the fruit within a sward canopy comprised of tendrilsin place of leaflets is discussed. However, there was stilla significant reduction in seed yield associated with the leaflessmutant despite a normal complement of ovule initials. The growthcurves and mean dry weights per seed were not significantlydifferent between phenotypes. In fruit of the conventional phenotypethe attainment of maximal pod wall weight coincided with theinitiation of seed fillingonday 15 after anthesis. In a seriesof isogenic lines the gene af delayed maximal wall developmentby 6 days whilst the gene st markedly lowered the maximum wallweight attainable. Possible causes of yield reduction in theleafless mutant are discussed in relation to the observed actionof these genes in the homozygous double recessive condition.     

Plasticity of reproductive components at different stages of development in the annual plant Thlaspi arvense L.

Summary This study examines the effect of different densities and the removal of all neighbours at different stages of development on all components of reproduction in the inbreeding annual Thlaspi arvense L. A 64-fold increase in density significantly reduced all repooductive components. The number of flower buds per plant was decreased most strongly; the order of decreasing plasticity among the other components was number of capsules per flower, individual seed weight, ovule number per capsule, flowers per flower bud and seeds per ovule. Removing neighbours at all stages of development increased seed yield of plants in comparison to the control without density reduction, but patterns of plasticity depended strongly on time of treatment. The main effect of the removal of neighbours at the vegetative stage was to increase the number of flowers per plant, but the number of ovules per capsule and seed weight increased also, and abortion of capsules decreased. Removing neighbours at the onset of flowering initially failed to affect flower number per plant, instead it resulted in a strong reduction of capsule abortion and an increase in seed weight. However, several weeks after flowering had initially ceased, fresh lateral inflorescences were produced, resulting in a second flush of reproduction. Removing neighbours at the stage of fruit ripening resulted at first only in an increase in seed witht, but later a second reproductive phase occurred. Fresh lateral branches were produced, but the apical meristem was also reactivated. The overall pattern of plasticity among all reproductive components in response to a removal of neighbours was the same as in response to density. The position of a capsule along the inflorescence influenced its number of ovules, the rate of seed abortion and the mean weight of seeds, with the type of effect depending on the developmental stage at which neighbours were removed. Significant negative correlations were found between the mean weight of seeds and the number of seeds in a capsule under all treatments.     

Flowering in Pisum: A Fifth Locus, Veg

In addition to the known loci, If, e, sn and hr, a fifth locus,veg, is shown to control flowering in peas. Regardless of thegenotype for the other flowering genes, plants homozygous forthe gene veg did not initiate flower buds under a wide rangeof photoperiod and temperature regimes, including those normallyhighly promotory in peas. Treatment with various plant growthsubstances and grafting to stocks known to promote floweringalso failed to cause initiation. Gene veg prevented expressionof allelic differences at the If locus but segregation for allelesat the sn and hr loci was clearly visible by examination ofseveral vegetative characteristics. For example, sn hr veg andSn hr veg plants showed an opening of the apical bud, productionof lateral branches, and a reduction in growth rate, leafletsize, internode length and stem thickness at approx the sametime as sn hr Veg and Sn hr Veg plants carrying the Lf allelecommenced fruit production, respectively. The graft-transmissibleinhibitor controlled by gene Sn is therefore not specific forthe transition from vegetative to reproductive growth. Geneveg allows the processes leading to apical and foliar senescenceto be examined independently of any effect of flowering andfruiting. We found that gene Sn influenced the total numberof leaves expanded in veg plants but not the time of shoot senescence,which, in plants without flowers and fruits appeared to resultfrom failure of the root system. Pisum sativum L., garden pea, flowering, senescence, genetics     

A Population Study to Aid the Selection of Improved Dried Pea (Pisum sativum) Crop Plants

The effect of genotype and plant density, over the range from100 to 277 plants m^–2, on plant to plant variation inprecision sown microplots has been assessed for three ‘leafless’(afafstst) pea (Pisum sativum) lines. This range of plantingdensities did not significantly affect the total above groundbiological yield per unit area of two of the genotypes (BS5and BS4) whereas the biological yield of the third (BS151) declinedat densities above 156 plants m^–2. The differences weredue to changes in seed yield. The effect of planting densityon the variation between plants for biological yield withinthe microplots differed between the genotypes. The distributionpattern of BS4 and BS5 changed from normal to skewed with increaseddensity, while the distribution for BS151 remained skewed atall planting densities. The differences between the three genotypes in the proportionof biological yield partitioned into seed yield (harvest index)on a unit area basis was due almost entirely to the differencesin structure of the plant populations. The maximum level ofpartitioning by individual plants was similar for all threelines. The difference between this maximum for an individualand the crop harvest index therefore represents the area forimprovement of crop harvest index through breeding. It is suggestedthat improvements in dried pea yields will come, therefore,by selecting plants which form more uniform populations withregard to plant size and to the proportion of plant biomasspartitioned into seed (plant harvest index). Pisum sativum, leafless pea, population, genetic variation, distribution patterns, harvest index     

Some Effects of Indol-3-yl Acetic Acid on Lateral Root Development in Attached and Excised Roots of Pisum sativum L.

The effect of continuous exposure to indol-3-yl acetic acid(IAA) on primordium initiation and their subsequent emergenceas lateral roots was determined in excised and attached rootsof Pisum sativum. IAA was found to stimulate the number of primordiainitiated per centimetre of attached or excised primary. Similarly,lateral emergence in terms of the number produced per centimetreof primary was promoted in the presence of IAA. This stimulationof lateral emergence even took place in excised roots whichwere 1 cm in length at the onset of culture and which neverproduced secondary roots over a 6-d culture period when grownin the absence of auxin. These effects of IAA on lateral rootdevelopment have been considered in relation to the concurrentchanges which take place in proliferative activity in the apicalmeristem of the primary root during exposure to auxin. Pisum sativum, garden pea, anlage, primordium, emerged lateral, cell proliferation, indol-3-yl acetic acid     

Factors affecting pollinator movement and plant fitness in a specialized pollination system

The rate of pollen exchange within and among flowers may depend on pollinator attraction traits such as floral display size and flowering plant density. Variations in these traits may influence pollinator movements, pollen receipt, and seed number. To assess how floral display size and flowering plant density affect parameters of pollinator visitation rate, pollen receipt per flower, seed number per fruit and the between-plant pollinator movements, we studied the self-incompatible plant, Nierembergia linariifolia. Per-flower pollinator visitation rate and bout length increased linearly with increasing floral display size. Pollen receipt per flower increased linearly with increasing flowering plant density. For seed number per fruit, a polynomial model describing an increased seed number per fruit at low density and a decreased seed number per fruit at high density provided a significant fit. Per-flower pollinator visitation rate was not associated with pollen receipt per flower and seed number per fruit. Bees visited plants located near to the center of the population more frequently than plants located at the periphery. Increases in both floral display size and flowering plant density led to an increased chance of a plant being chosen as the center of the pollinator foraging area. These results suggest that even though large floral displays and high flowering plant density are traits that attract more pollinators, they may also reduce potential mate diversity by restricting pollen movement to conspecific mates that are closely located.     

EFFECT OF DEFOLIATION ON GENDER EXPRESSION AND FRUIT SET IN PASSIFLORA INCARNATA

We examined the effect of defoliation on gender expression and fruit set in a north-central Florida population of the andromonoecious vine, Passiflora incarnata, during the 1984 flowering season. At three times during the flowering season (May, June, July), the leaves adjacent to flowers at four stages of development were removed, and subsequent sex and fate of each flower were determined. Defoliation affected gender of the flower by significantly decreasing the probability that the styles of the flower would deflex and the flower would thus function only as a pollen donor. Flowers were sensitive to defoliation at any stage before anthesis, though the sensitivity appeared to decrease in the most mature category of flower buds. Fruit set of hermaphroditic flowers remained unaffected by defoliation, but the probability of fruit set and gender expression were significantly influenced by the time of the flowering season. We conclude that the local photosynthate environment determines flower gender in Passiflora, but branch or entire plant photosynthate resources can compensate for local resource fluctuations and play an important role in fruit set and flower bud abortion. The sexual lability of Passiflora incarnata appears to be an adaptation to uncertain resource levels at a fine scale, caused by nearby developing fruits and the possibility of herbivore defoliation.     

Carbon Transfer and Partitioning between Vegetative and Reproductive Organs in Pisum sativum L

Assimilate partitioning was studied in the common pea (Pisum sativum L.) by feeding ¹⁴CO₂ to whole plants and measuring radioactivity in different organs 48 hours after labeling. Two experimental protocols were used. For the first, one reproductive node was darkened with an aluminum foil, to prevent photosynthesis during labeling. The aim was to study assimilate translocation among nodes. The second was carried out to assess any priority among sinks. Whole plants were shaded, during labeling, to reduce carbon assimilation. Various developmental stages between the onset of flowering and the final stage in seed abortion of the last pod were chosen for labeling. When all photosynthetic structures at the first reproductive node were darkened at any stage of development after the formation of the first flower, the first pod was supplied with assimilates from other nodes. In contrast, later developed pods, when photosynthetic structures at their node were darkened, received assimilates from other nodes only when they were beyond their final stage in seed abortion. Reducing illumination to 30% did not change distribution of assimilated carbon between vegetative and reproductive structures, nor among pods. It appears that the relative proportion of ¹⁴C allocated to any one pod, compared to other pods, depends on the dry weight of that pod as a proportion of the total reproductive dry weight. When the plant was growing actively, following the start of the reproductive phase until a few days before the end of flowering, the top of the plant (i.e., all the organs above the last opened flower) had a higher sink strength and a higher relative specific activity than pods, suggesting that it was a more competitive sink for assimilates. The pattern of assimilate distribution described here provides an explanation for pod and seed abortion.     

The Effect of Source-Sink Alterations on Soybean Seed Growth

Soybeans (Glycine max L. Merrill) were grown in the greenhouseand in the field to investigate the effect of variations inthe assimilate supply during the linear phase of seed developmenton the rate and duration of growth of individual seeds. Increasedassimilate supplies, created by partial fruit removal, increasedrates of dry matter accumulation, duration of seed growth, andfinal seed size (weight per seed). Reductions in the supplyof assimilate to the developing seed, created by shading (60per cent) the plants during the linear phase of seed development,lowered seed growth rate but did not affect final seed sizebecause of a longer duration of seed growth. Nitrogen stressduring seed development, created by removing N from the nutrientmedium, did not affect seed growth rate but shortened the durationof seed growth and reduced final seed size. The data indicatethat the growth characteristics of soybean seed are influencedby the supply of assimilate to the seed during the linear phaseof seed development. Glycine max L., soybean, seed growth rate, duration of seed growth, effective filling period