Phenotypic correlations among plant parts in Iberian Papilionoideae (Fabaceae)

BACKGROUND AND AIMS: Different plant organs may show varying degrees of form diversification or conservatism across phylogenetically related taxa. The present study uses data from a recent systematic study of Iberian Papilionoideae to investigate diversification and covariation in reproductive and vegetative plant parts. The appropriateness of imprecise (but comprehensive) taxonomic quantitative information is tested. METHODS: Organ size covariation and phenotypic correlations were studied among tribes, genera and species. Scale relationships were investigated by Reduced Major Axis regression. Variables used were the maximum dimensions of calyx, corolla, keel petal, fruit, seed, stipule, leaflet and petiole. KEY RESULTS: As regards tribe averages, the length of the corolla and that of calyx correlated positively and significantly. In contrast, pod length was unrelated to corolla size and largely tribe-specific. Within genera, the sizes of calyx, corolla and fruit sometimes covaried linearly (e.g. Lathyrus species) and other times did not (Genista, Astragalus). CONCLUSIONS: Information from taxonomic studies can be useful to establish major phenotypic correlations in plants. Results underscore the implications of tribal ownership in the Papilionoideae and illustrate the extensive morphological diversification of pods relative to flowers in this group.     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

ECOLOGICAL CONSEQUENCES AND PHENOTYPIC CORRELATES OF PETAL SIZE VARIATION IN WILD RADISH,RAPHANUS SATIVUS (BRASSICACEAE)

In this paper we examine some ecological consequences and phenotypic correlates of flower size variation in wild radish, Raphanus sativus. Mean corolla diameter varied significantly among individuals within natural populations of R. sativus in California. On the average, almost 40% of flower biomass was allocated to corolla tissue. In field experiments, pollinator visitation increased significantly with corolla size. Large flowers also accumulated more nectar when pollinators were excluded from plants. In three populations, corolla size was positively correlated with allocation to pollen per flower (either anther weight or pollen grain number), but there was usually no phenotypic relationship between corolla size and several measures of female allocation (ovule number per flower, proportion fruit set, and total seed mass per fruit). Plants growing in the field produced fewer large flowers per unit of stem, and stem biomass was negatively related to corolla size for plants grown under controlled greenhouse conditions. Male and female fitness may covary differently with allocation to attractive floral features in species such as R. sativus, where seed production is often limited by resources rather than by pollen.     

Effect of ovule position within the pod on the probability of seed production in Bauhinia ungulata (Fabaceae)

BACKGROUND AND AIMS: It has been claimed that ovules linearly ordered within a fruit differ in their probabilities of reaching maturity. This was investigated by studying the effect the position of an ovule within the pod has on seed abortion and seed production in Bauhinia ungulata. METHODS: Fruits collected during the dry seasons of 1999, 2000 and 2001 were opened, and the number, position and status of each ovule within the fruit were recorded. A GLM model was used to assess the effects of population, tree identity and ovule position within the pod on ovule fertilization, seed abortion, seed damage and seed maturation in two populations of B. ungulata. KEY RESULTS: Nearly 30% of the ovules were not fertilized in 1999; this percentage dropped to 5% the following two years. Seed abortion (50%) and seed damage (15%) were the same every year during the study period. Only 15% of the initial ovules developed into mature seeds in 1999; this value increased to 35% in 2000 and 2001. However, seed survivorship was dependent on the position of the ovule within the pod; non-fertilized and early aborted ovules were found more often near the basal end of the ovary. The frequency of seed damage was not affected by position. Mature seeds were found mainly in the stylar half of fruits, where ovules are likely to be fertilized by fast pollen tubes. CONCLUSIONS: The pattern of seed production in B. ungulata is non-random but is dependent upon the position of the ovule within the pod. The results suggest that the seeds produced within a fruit might differ in their vigour.     

药物植物桃儿七不同种群种子产量初步研究

对药用植物桃儿七5个不同种群种子产量的研究表明：各种群中每花平均胚珠数、每果实平均种子数及种群平均结实率与种群所在的海拔高度呈正相关;不同种群个体平均胚珠数的变异范围为58.97-87.97,平均种子数变异范围为40.02-80.58,平均结籽率变异范围为61.29%-91.60%;果料内单粒种子平均干重与种群分布海拔呈明显负相关,与果实内种子数呈弱负相关;同一种群在不同年份间平均各上子产量差异不大;种子产量除主要受个体营养状况影响外,还受海拔及其它因素的影响;果实内种子数与果实大小及重量呈正相关,但单粒种子平均重量与果实大小及果实内种子总重量无显著相关。     

Morphology, anatomy and histochemistry of Salicornioideae (Chenopodiaceae) fruits and seeds

BACKGROUND AND AIMS: The subfamily Salicornioideae (Chenopodiaceae) are a taxonomically difficult group largely due to the lack of diagnostic characters available to delineate tribal- and generic-level boundaries; a consequence of their reduced floral and vegetative features. This study examined the variation in fruits and seeds across both tribes of the Salicornioideae to assess if characters support traditional taxonomic sections. METHODS: Light microscopy, environmental scanning electron microscopy and anatomical ultra-thin sectioning were employed to examine variation in fruits and seeds. Sixty-eight representatives across 14 of the 15 genera currently recognized within the tribes Halopeplideae and Salicornieae were examined to determine whether characters support current taxonomic groups. KEY RESULTS: Characters such as seed coat structure, embryo shape, seed orientation, the forms of seed storage proteins and carbohydrates show variation within the Salicornioideae and may be phylogenetically useful. The campylotropous ovule typical of the Chenopodiaceae generally results in a curved embryo; however, many Halosarcia and Sclerostegia species have straight embryos and in Salicornia and Sarcocornia the large peripheral embryo appears bent rather than curved. Seed coat ornamentation of Microcnemum and Arthrocnemum is distinct from other Salicornioideae as the elongated epidermal cells of the exotesta have convex walls. Histochemical stains of anatomical sections of cotyledon cells showed protein bodies were variable in shape, and starch grains were present in some species, namely Salicornia bigelovii, S. europaea and Allenrolfea occidentalis. CONCLUSIONS: While fruits and seeds were found to be variable within the subfamily, no synapomorphic characters support the tribe Halopeplideae as these genera have crustaceous seed coats, curved embryos and abundant perisperm; features characteristic of many of the tribe Salicornieae. The endemic Australian genera are closely related and few seed and fruit characters are diagnostic at the generic level. Nineteen characters identified as being potentially informative will be included in future phylogenetic analyses of the subfamily.     

Seed set and germination in some wild species of Vicia from SW Europe (Spain)

We studied seed and ovule production, predation and abortion in 14 taxa of the genus Vicia. Germination capacity, seed vigour and germination kinetics after scarification were also evaluated, as well as the influence of light on these parameters. Pod size correlated with the number and weight of viable seeds, and with the total number of ovules and the number of undeveloped ovules per pod. Species with smaller seeds had lower rates of abortion than those with larger seeds, and seedling size correlated strongly with seed weight in this genus. In most taxa seed hardness was not great, and the scarifying agent used—sulfuric acid with different times of exposure—led in some cases to 100% germination. Light conditions did not notably influence germination. Germination speed was favored by scarification under both dark and light conditions, and vigour index was greater in scarified than in control seeds. In most taxa, germination after scarification fit an exponential curve, with the exception of Vicia lutea ssp. vestita, which showed very slow velocity.     

Correlations and heritability of the characters determining the seed yield of the long-raceme alfalfa (Medicago sativa L.)

Phenotypic correlation coefficients and heritability of the characters controlling seed yield of long-raceme forms of alfalfa was determined. It was found that seed yield per plant, which was positively correlated with 10 out of 12 analysed characters, depended upon the number of pods per raceme and the number of seeds per pod. Variability of these characters determined about 60% of the variability of seed yield. Multiple linear regression and phenotypic correlations show that simultaneous selection for increased pod number per raceme and increased seed number per pod and raceme length resulted in enhanced seed yield potential. The share of the additive genetic effects in the phenotypic variance for number of pods per raceme was low and about 21-23%, while for number of seeds per pod and per raceme amounted to about 50%. The expected genetic progress in recombination breeding for number of seeds per pod and number of seeds per raceme will be of medium magnitude, while one cannot expect any rapid and considerable progress in the number of pods per raceme. Considering the high positive correlation between raceme length and number of pods and seeds per raceme, one should conclude that raceme length can be an important criterion in selection of plants showing a high seed productivity.     

中国木蓝属两个分布新记录

报道并描述了采自云南南部西双版纳具有1小叶的新分布种美脉木蓝Indigofera caloneura Kurz和采自广东具有单叶的新分布种心叶木蓝I. cordifolia Heyne ex Roth.。在中国还有另外3个具有单叶的种类, 即单叶木蓝I. linifolia、刺荚木蓝I. nummularifolia和远志木蓝I. squalida。通过从植物习性、茎、叶的形态与大小及毛被、托叶、花序、花冠形状与毛被、花药、子房毛被、果实的形态与大小、所含种子数目等特征的显微观察, 比较了国产单叶类木蓝属Indigofera植物的形态学特征差异, 并针对茎、叶、毛被、花冠旗瓣和果实形状等5个最重要的特征进行了讨论。     

Patterns of fruit and seed production inBauhinia ungulata (Leguminosae)

Patterns of seed and fruit production ofBauhinia ungulata, a small tree legume indigenous in tropical America, were studied in Costa Rica. Only about 8% of flowers produced fruits. The average pod had 19 ovules and about two thirds of these began seed development, with mature pods containing an average of 9.7 mature undamaged seeds. About half of the mature pods were damaged by herbivores and within these, 27% of ovules or seeds had been eaten. Among trees there was no significant variation in pod production, but the number of ovules per pod and seed production per pod varied significantly. Within infructescences most pods were retained at middle positions. Within pods, the probability of an ovule developing into a seed increased toward the distal end. The pattern of seed and fruit production in this species agrees well in general with that reported for other neotropical legumes. The abortion of seeds and fruits can be regarded as a way of controlling maternal investment, and as a response to herbivory.     

Variation in ovule and seed size and associated size-number trade-offs in angiosperms

Unlike pollen and seed size, the extent and causes of variation in ovule size remain unexplored. Based on 45 angiosperm species, we assessed whether intra- and interspecific variation in ovule size is consistent with cost minimization during ovule production or allows maternal plants to dominate conflict with their seeds concerning resource investment. Despite considerable intraspecific variation in ovule volume (mean CV = 0.356), ovule production by few species was subject to a size-number trade-off. Among the sampled species, ovule volume varied two orders of magnitude, whereas seed volume varied four orders of magnitude. Ovule volume varied positively among species with flower mass and negatively with ovule number. Tenuinucellate ovules were generally larger that crassinucellate ovules, and species with apical placentation (which mostly have uniovulate ovaries) had smaller ovules than those with other placentation types. Seed volume varied positively among species with fruit mass and seed development time, but negatively with seed number. Seeds grew a median 93-fold larger than the ovules from which they originated. Our results provide equivocal evidence that selection minimizes ovule size to allow efficient resource allocation after fertilization, but stronger evidence that ovule size affords maternal plants an advantage in parent-offspring conflict.     

The influence of plant and corolla size on pollen deposition and seed set in Geranium sanguineum (Geraniaceae)

In a coastal grassland area the influence of plant and corolla size on insect visitation rate, pollen deposition on stigmas and seed set was measured in a natural population of Geranium sanguineum. Visits by bumble bees, the primal pollinators, were significantly affected by plant size measured as the number of flowers, and differences in visit rates among plants were explained by size differences among these. On average, larger plants had bigger corollas. The mean corolla area affected pollen deposition positively whereas deposition did neither affect seed set (number per flower) nor seed weight. Insects reaction to corolla size was dependent on plant size, and thus we found the effect of corolla size on pollen deposition to decline with plant size. Instead pollen deposition per plant seemed to be correlated to overall floral display. We argued that resource status of the individual plants could explain corolla size and seed set variation. The finding of some plant setting seeds while other plant with comparable size did not set any seeds could not be explained satisfactory by resource limitation.     

Seed abortion in wind-dispersed pods of Dalbergia sissoo: maternal regulation or sibling rivalry?

Summary Dalbergia sissoo, a wind-dispersed tropical tree, shows a positively skewed distribution of seeds per pod. This is attributed to the enhanced dispersal advantage of few-seeded pods due to their reduced wing loading (ratio of weight to pod surface area) and low settling velocity. The proximate mechanisms causing the positively skewed distribution were investigated. The distribution could not be attributed to the distribution pattern of ovule number per ovary, pollen grain limitation, lack of ovule fertilization, or post-fertilization elimination of many-seeded pods. Rather, it was caused by the post-fertilization abortion of seeds within a pod 2 weeks after fertilization. This intra-pod seed abortion (IPSA) is due to a dominance hierarchy of fertilized ovules from the distal (near stigma) to the basal end, generated by the temporal differences in fertilization. The dominant developing seeds at the distal end cause the abortion of others through the production and diffusion of an aborting agent. When the dominance hierarchy of the siblings is not intense, pods are formed with more than one seed. We argue that the positively skewed distribution of seeds per pod is not due to maternal regulation but is a result of sibling rivalry. We propose that this sibling rivalry is generated by genetic differences in pollen grain fitness and disucss the results in the context of parent-offspring conflict.     

MEASUREMENTS OF NATURAL SELECTION ON FLORAL TRAITS IN WILD RADISH (RAPHANUS RAPHANISTRUM). I. SELECTION THROUGH LIFETIME FEMALE FITNESS

Although the role of natural selection in the evolution of floral traits has been of great interest to biologists since Darwin, studies of selection on floral traits through differences in lifetime fitness have been rare. We measured selection acting on flower number, flower size, stigma exsertion, and ovule number per flower using field data on lifetime female fitness (seed production) in wild radish, Raphanus raphanistrum. The patterns of selection were reasonably consistent across three field seasons, with strong directional selection for increased flower production in all three years, weaker selection for increased ovule number per flower in two years, and selection for increased flower size in one year. The causes of the selection were investigated using path analysis combined with multiplicative fitness components. Increased flower production increased fruit production directly, and increased numbers of ovules per flower increased the number of seeds per fruit in all three years; pollinator visitation did not influence either of these fitness components. Increased flower size was associated with increases in both the number of fruit and the number of seeds per fruit in one year, with the latter relationship being stronger. Total lifetime seed production was affected more strongly by differences in fruit production than by differences in either the number of seeds per fruit or the proportion of fertilized seeds that were viable, but all three fitness components were positively correlated with total seed production.     

Ovule Development and Determination of Seed Number Per Pod in Oilseed Rape (Brassica napus L.)

Seed number per pod at maturity over the terminal raceme ofsingle plants of oilseed rape is closely correlated to the percentageof ovules with complete embryo sacs (ovule fertility) at floweropening. Approximately one-third of the ovules did not containan embryo sac and sterility, due to the absence of embryo sac,accounted for most of the difference between the numbers ofovules and seeds. Within the terminal raceme, both a decreasedproportion of fertile ovules and a lower number of ovules perovary in apical flowers contributed to the lower number of seedsper pod in the mature apical pods compared to the basal ones.A study of ovule development before flower opening showed thatdifferences in the differentiation of the embryo sacs arosebefore the buds were 40 mm long and probably involved the stagesof meiosis II and/or differentiation of the chalazal megaspore. Key words: Oilseed rape, ovule development, seed number per pod     

Long-term variation in seed mass and seed production in populations of Paris quadrifolia

Seed production of the perennial herb Paris quadrifolia L. (Liliaceae) was investigated in five populations in northern Poland. The long-term seed production per square metre differed significantly among populations and years. Moreover, throughout the 7-year study period, 30% of both whole ripe fruits and seeds alone were predated. Variation in seed mass per fruit in both space and time was significant. Throughout the 7-year study, nearly all the marked individuals produced fruits once every 2 years. The most frequent break between fruiting was 2 years and the longest was 5 years. Only the mean seed mass in fruits of the same individual varied significantly over subsequent years. In the five populations, the number of ovules, number of seeds in the fruit and seed mass varied significantly between populations. However seed/ovule ratio did not differ in fruits in the five populations. The seed mass/number trade-off in fruits was strongly partially correlated when the effect of total seed mass was considered. Breeding experiments suggest that P. quadrifolia has a substantial capacity for both inbreeding and outbreeding. There were no significant differences in the seed/ovule ratio, seed number or seed mass in fruits produced from bagged or control flowers. However, both seed/ovule ratio and number of seeds were significantly lower in fruits from emasculated flowers.     

Sex allocation and mating systems in pigeonpea,Cajanus cajan (Fabaceae)

Sex allocation theory predicts that: (1) resources allocated to androecium should decrease with an increase in selfing, (2) a decrease in androecium biomass should be accompanied by an increase in the biomass of pistils, and (3) a decrease in androecium biomass should be coupled with a decrease in flower size, specifically corolla biomass. Another predicted change in reproductive traits associated with variation in selfing concerns seed to ovule ratios, but does not directly stem from sex allocation theory. It has been postulated that seed to ovule ratios should be positively correlated with the amount of selfing. These predictions were tested for six accessions of pigeonpea,Cajanus cajan L., that differed in selfing rates. The results were remarkably in accordance with the predictions. We conclude that sex allocation theory provides a powerful tool to understand the evolution of many reproductive traits in plants.     

SEED SIZE AND SEED PACKAGING VARIATION IN BAPTISIA LANCEOLATA (FABACEAE)

Pods and seeds of field-collected Baptisia lanceolala plants were analyzed to partition seed weight and seed packaging trait variance among and within plants and to detect relationships between these traits. Packaging traits studied were: pod weight, seed weight per pod, number of seeds per pod, mean weight of seeds per pod, proportion seed weight of total pod weight, and pod weight per seed. Significant among-plant variation was found for seed weight and all packaging traits. Within plants, positive correlations were found between number of seeds per pod and pod dry weight and between the proportion seed weight of total pod weight and number of seeds per pod. Pod weight per seed was negatively correlated with number of seeds per pod. Most plants had a negative correlation between mean seed weight and number of seeds per pod. When compared with an equality of slopes test, slopes of regressions of the above pairs of traits were found to differ among plants. Among plants, the same relationships were found, except for the latter two traits, which were not correlated. These within-plant patterns may represent constraints on seed weight variance imposed by the seed package. This view is supported by a positive correlation between packaging trait variance and seed weight variance. Packaging-related constraints could have an effect on seed weight in this and other species. If these phenotypic constraints have a genetic basis, then selection on seed packaging could change seed weight in a way different from that which might be predicted by considering seed weight alone.     

Factors Influencing Seed: Ovule Ratios and Reproductive Success in Four Cleistogamous Species: A Comparison between Two Flower Types

Abstract: Ovule and seed numbers, and the ratio between the two, were determined in chasmogamous (CH) and cleistogamous (CL) flowers/fruits in four perennial herbs: Viola hirta, V. mirabilis, V. riviniana, and Oxalis acetosella. The results were applied to data from earlier studies on fruit set and progeny performance in these species, to find implications for factors influencing seed: ovule (S: O) ratio, to relate S: O ratio to other fitness components, and to compare final numbers of seedlings per initiated ovule between the two reproductive modes. Mechanisms considered to have a potential impact on S: O ratios were pollen limitation, resource limitation, and "genetic load". In V. mirabilis and O. acetosella, mean S: O ratio was significantly higher in the CL phase than in the CH phase; in V. hirta and V. riviniana, there was no difference. Pollen limitation seemed improbable as a factor important for seed set in all species, and no indication was found of resource limitation. There was a negative correlation between S: O ratio and seed germination rate in O. acetosella, indicating that seed abortion, particularly in CH fruits, may be a way to purge inferior zygotes. The high CH S: O ratios in all species suggest that substantial selfing also occurs in this phase. The final number of seedlings per ovule did not differ significantly between modes in any of the species, partly due to poorer germination in CL seeds. This indicates that, despite the often lower pre-emergent reproductive success (fruit and seed set) in the CH phase, total reproductive success will not differ between the two modes because post-emergent selection is stronger on CL progeny. The strongest limitation on pre-emergent reproductive success in both phases is apparently related to flower/fruit abortion.     

海拔对青藏高原东部毛茛科植物繁殖特征和资源分配的影响

通过野外采样对青藏高原东部高寒草甸上两个海拔间3种常见毛茛科植物条裂银莲花(Anemone trullifo-liavar.linearis)、粗距翠雀花(Delphinium pachycentrumHemsl.)和钝裂银莲花(Anemone obtusiloba)的繁殖性状和资源分配进行了研究,并对植物在极端环境下采取的繁殖策略及繁殖模式进行了探讨.结果显示:(1)3个物种的株高都随着海拔的升高而降低;同一物种的花大小在不同海拔间均无显著差异;条裂银莲花单个种子重随海拔的升高而增加,但虫食数/株、结籽率和种子数/株均随着海拔的升高而减少;钝裂银莲花的繁殖分配、虫食数/株、种子数/株和单个种子重均随着海拔的升高而减小.(2)不同海拔条裂银莲花的个体大小与单花重、雄蕊重、虫食数/株均呈显著正相关,且回归斜率在海拔间有显著差异.粗距翠雀花花期的雄蕊重与个体大小呈显著正相关,但这种异速关系不受海拔和个体大小的影响;钝裂银莲花的个体大小与花期的所有繁殖特征以及果期的种子重/果实均呈显著正相关,不同海拔间个体大小与种子重/果实的斜率差异显著,且与花期各繁殖性状异速关系的截距差异显著.研究表明海拔对植物的个体大小以及种群间的繁殖对策和繁殖成功率有着重要的影响.