The inheritance of partial self-compatibility in Brassica oleracea L. inbreds homozygous for different S-alleles

Summary In a study of partial self-compatibility in Brassica oleracea, flower number, seeded siliqua and seed production were recorded on self and cross-pollinated inflorescences of 32 progenies obtained by inter-crossing and selfing 8 plants homozygous for the incompatibility alleles S₂, S₅, S₁₅ and S₄₅.Progeny differences for both self-and out-cross seed production could be largely attributed to G.C.A. effects which were essentially uncorrelated. For cross-pollinated inflorescences heterosis was also important. Significant differences were found for selfed seed set and its two components, the proportion of flowers producing seeded siliquae and the numbers of seed per seeded siliqua, between parents with the same S-allele which could not be attributed to S-genotype alone. No evidence of increased self-compatibility in particular S-allele heterozygotes (mutual weakening) could be found.Outcross seed production depended primarily on the numbers of seeds set per seeded siliqua while self seed production was largely determined by the proportion of flowers which produced seeded siliqua. It is suggested that this is a key character for the production of inbred lines with reduced partial self-compatibility.     

Ovule pre-emption and pollen limitation in a self-fertile perennial herb (Blandfordia grandiflora,Liliaceae)

The extent, frequency and causes of pollenlimited seed production were examined in partially selffertile populations of Blandfordia grandiflora for 2 years. Percentage seed set of open-pollinated plants (50–57%) did not differ within or between years, and was about 19% less than experimentally cross-pollinated plants (70–75%). Floral visits by honeybees did not differ through the flowering season and the number of pollen grains deposited on stigmas within 12 h of flowers opening exceeded the number of ovules per flower, indicating that the quality rather than the quantity of pollination limited seed set. Pollen limitation was caused by concurrent self- and cross-pollination and the subsequent abortion of some selfed ovules due to inbreeding depression. Natural seed set (55%) was intermediate between selfed (43%) and crossed (75%) flowers and was not increased when flowers that had been available to pollinators for 24 h were hand cross-pollinated, suggesting that ovules were already fertilized. Similarly, experimental pollination with both cross and self pollen within 24 h of flowers opening did not increase seed set relative to natural seed set, indicating that both cross- and self-fertilizations had occurred. In contrast, when selfing followed crossing by 48 h, or vice versa, seed set did not differ from crossed-only or selfed-only flowers, respectively, indicating that ovules were pre-empted by the first pollination. Collectively, these results indicate that under natural conditions self pollen pre-empts ovules, rendering them unavailable for cross-fertilization. This selfing reduces fecundity by 50%, as estimated from the natural production of cross seeds when selfing was prevented. Consequently, selection should favour floral traits, such as increased stigma-anther separation or protandry, that reduce interference between male and female functions that leads to selfing.     

Pollination ecology of Penstemon roseus (Plantaginaceae), an endemic perennial shifted toward hummingbird specialization?

Summary Pink-flowered tubular Penstemon roseus (Plantaginaceae), which has shifted partially to hummingbird pollination, blooms on high-elevation slopes in the mountains in Tlaxcala, Mexico. We studied the interactions between pollinator visitation rates to flowers, pollen removal and deposition, flower size, and nectar removal frequency on seed production in P. roseus. We combine observational and experimental studies in two contrasting natural populations. Our manual pollinations revealed that P. roseus is fully self-compatible. Autonomous self- and manual self-pollinated flowers matured as many seeds as when outcrossed, but outcrossing seems to become better than selfing as the flowering season progressed. Early in the season flowers that were bagged and hand-selfed, hand-outcrossed, or autonomously selfed, or unbagged and naturally pollinated had equal seed set in all four treatments. But later in the season, outcross pollen gave approximately twice as much seed set as the two self-treatments. Low levels of pollen receipt and pollen removal were consistent with the long time elapsed for a given plant to be visited by hummingbirds, which suggests pollen shortage in both sites. Despite differences in pollinator visitation rates to flowers, probability of flower visitation, removal and deposition of pollen, and nectar production rates between populations, we found that total nectar production had no effect on seed production at either site. The daily nectar secretion rate of 0.3–0.65 mg sugar per flower per 1–3 days was low relative to other hummingbird-adapted Penstemon species (typical range: 1.5–5 mg sugar per flower), and it might be intermediate between hummingbird- and bee-adapted Penstemon flowers. Our results support the hypothesis about a shift toward hummingbird pollination, and provide an example of a ‘despecialized’ Penstemon species, which attracts high-energy pollinators (hummingbirds) and profits from outcrossing, but retains bee-syndrome floral traits and low sugar production rates.     

FACTORS LIMITING FRUIT AND SEED PRODUCTION OF A TEMPERATE SHRUB,STAPHYLEA TRIFOLIA L. (STAPHYLEACEAE)

Staphylea trifolia L., the bladdernut, is a self-incompatible temperate woodland shrub that flowers in May in Illinois. Factors limiting reproduction were studied at four levels: 1) Seeds/fruit. Seed production in open-pollinated fruit was frequently limited by too few fertilized ovules. Seed production in hand-cross-pollinated fruit was limited by resources or dispersal constraints: seed abortion rates were higher in hand-cross-pollinated fruits than in open-pollinated fruits. 2) Fruits/flower and 3) fruits/inflorescence. The number of fruits set and matured per flower and per infloresence in the open-pollinated treatment was limited by the number of flowers naturally cross-pollinated. In hand-cross-pollinated inflorescences, fruit set was not limited by resources even though fruit set was ten times greater than in the open-pollinated treatment. Evidence that resources limited fruit maturation in the hand-cross-pollinated inflorescences was equivocal. In hand-cross-pollinated flowers, fruit set was lowest when cold nights followed pollination, suggesting that cool temperatures limited postpollination physiological processes. 4) Fruits/individual. Early-flowering individuals matured fewer fruits than later-flowering individuals. Within the latter group, fruit production increased with plant size, although a relatively small individual matured the maximum number of fruits. Flowering phenologies and size of individuals varied among patches, resulting in differential reproductive success of patches.     

Intra-inflorescence variation in floral traits and reproductive success of the hermaphrodite Silene acutifolia

BACKGROUND AND AIMS: Intraspecific variation in floral components and reproductive success is often located at the intra-individual level. The arrangement of flowers within inflorescences may explain a great deal of this variation. The variation in number of ovules, fruit set, number of seeds per fruit, seed set, seed weight and seed germination is investigated at different positions within the inflorescence of Silene acutifolia. METHODS: Data were obtained in natural populations, and germination experiments were conducted in a germination chamber. KEY RESULTS: The number of ovules, fruit set, number of seeds, seed set and seed weight, showed a significant decline from early (primary) position to later (tertiary) position. The patterns of intra-inflorescence variation were consistent in different populations and years of study. Seed germination showed an opposite pattern, seeds from primary position showed the lowest germination percentages and seeds from tertiary position the highest, although the effect of position on germination was only marginally significant. There was significant among-population variation in number of ovules per flower. Fruit set also varied significantly among populations, with lower fruit set in the smaller and more isolated population. No significant among-population differences were detected in number of seeds per fruit and seed set. Seeds from the smallest and more isolated population (Arnado) were the lightest. Seed germination showed strong differences between populations, seeds from Arnado started to germinate later, and showed the lowest final germination percentages. CONCLUSIONS: Architectural effects or resource competition are the most commonly proposed hypothesis to explain these patterns. Data suggest that there is less pollen available to pollinate tertiary flowers, and that there is not enough outcross pollen in Arnado. The germination percentages suggest that there is variation in the source of pollen within inflorescences, with high probability of receiving outcross pollen in flowers from primary position, and higher probability of geitonogamous crosses in tertiary flowers.     

Effect of ‘Mixtalol’ on growth,yield and yield components of Indian mustard (Brassica juncea)

Mixtalol (a mixture of long chain aliphatic alcohols varying in chain length from C₂₄ to C₃₂) applied to Brassica juncea plants as foliar spray caused an increase in secondary and tertiary branching with consequent enhancement in seed yield through increased number of inflorescences and siliquae per plant. The percentage of immature siliquae and shattering of siliquae decreased with this treatment. Mixtalol increased total dry matter of plants, partitioning coefficient and harvest index. The contents of starch, protein and oil were also higher in seeds from Mixtalol treated plants.     

Evidence for selective fruit abortion in Banksia spinulosa (Proteaceae)

Fruit set is consistently low in the genus Banksia (Proteaceae). One explanation for low fruit:flower ratios is that excess flowers allow plants to selectively abort inferior progeny thereby increasing the average quality of the seed crop. We examined whether self-pollinated fruits were aborted selectively in a partially self-compatible population of Banksia spinulosa. Inflorescences were divided in half and pollinated with cross pollen on both sides, self pollen on both sides or cross pollen on one side and self pollen on the other. Seed production was reduced significantly by 38% on fully self-pollinated compared to fully cross-pollinated inflorescences, indicating partial self-incompatibility or inbreeding depression. On inflorescences pollinated with both cross and self pollen, selfed seed set was reduced by 63% compared to crossed seed set. On the selfed half of these inflorescences, fewer fruits and seeds were produced, and more fruits had aborted seeds than on randomly selected halves of fully self-pollinated inflorescences. These results suggest that self-fertilized embryos were developed in the absence of outcrossed progeny but were selectively aborted when outcrossed progeny were present on inflorescences. To determine the amount of outcrossing occurring in the study population, outcrossing rates (t) of naturally pollinated plants were examined by electrophoretic assessment of progeny arrays. Values of I did not differ significantly from one, indicating complete outcrossing. High outcrossing, despite the high probability of self pollen receipt under natural conditions, is consistent with the conclusion that selective abortion occurs in B. spinulosa.     

Temporal patterns of seed set and decelerating fitness returns on female allocation in Zigadenus paniculatus (Liliaceae), an andromonoecious lily

I describe temporal patterns of seed production in the andromonoecious lily Zigadenus particulatus. Fruit set per flower and seed set per fruit declined through time within plants. Hand pollination experiments showed that this was not due to increasing pollen limitation. Nutrient supplementation had little effect on seed output, but leaf clipping reduced seed production, especially in late-blooming flowers, and removal of early-blooming flowers increased seed set by later flowers. Thus, the temporal pattern of seed output was due to declining availability of photosynthates. Plants with larger bulbs produced larger inflorescences, a greater proportion of hermaphrodite flowers, more fruits per hermaphrodite flower, and more seeds per fruit, but lost a greater fraction of their initial bulb mass as a consequence of fruiting. After controlling for the effects of bulb mass, plants with larger inflorescences produced a greater proportion of male flowers, and plants with more hermaphrodite flowers produced fewer fruits per hermaphrodite flower and fewer seeds per fruit. Thus, the female fitness gain curve was decelerating. The temporal decline in seed output provides a partial explanation for the parallel decline in allocation to pistils. However, a complete explanation for the pattern of gamete packaging requires an understanding of factors controlling male, as well as female, fitness.     

Phenotypic gender variation within inflorescences of the protogynous species Helleborus foetidus L. (Ranunculaceae)

In the protogynous species H. foetidus, I investigated if investment in sexual structures and seed set shows any trend with position in the flower-opening sequence. In four-flower inflorescences, stamen number decreased with flower position i.e. was highest for the earliest flower and lowest for the latest flower. Ovule number was significantly higher in the latest flower. Stamen and ovule number did not covary, indicating that there is no structural gender trade-off in this species. Seed set (i.e. percentage of ovules producing seed) did not differ between control plants and pollen-supplemented plants and the effect of pollen supplementation did not vary among positions. Seed set showed marginally significant differences between control plants and simulated-herbivory plants, but the effects of this treatment varied significantly among positions. Significant among-position variation in seed set was observed in the control plants but not the simulated-herbivory plants. Seed set of latest-opening flowers of simulated-herbivory plants was significantly higher than that of latest-opening flowers of control plants. H. foetidus support Brunet and Charlesworth's (1995) prediction that in plants with protogynous flowers, later-opening flowers should specialize as female flowers, at least under conditions of high resource availability.     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Pollen limitation and seed set associated with year-to-year variation in flowering of Gmelina arborea in a natural tropical forest

The year-to-year variations in flowering, pollen and fruit production in ten Gmelina arborea individuals in a natural forest were observed over a period of five years, 2009?2013, and pollinator visits were observed closely over two years, 2011 and 2012. A pollen supplementation experiment was also undertaken in two years (2011 and 2012) of contrasting flowering levels. Considerable year-to-year variations were observed in flower, pollen and fruit production. The observed variation represented a normal sequence of good-year and poor-year cycling which is most parsimoniously attributed to resource allocation. The average level of pollen production per individual tree in good flowering years oscillated between 6.6 and 9.62 × 10⁸, which in the poor flowering years was between 1.92 and 3.07 × 10⁸. The magnitude of pollen limitation across years was highly significant (p < 0.0001). Fruit set through supplemental pollination was 73% and 96% greater than that of the open-pollinated branches in the year 2011 and 2012, respectively. Pollen limitation, irrespective of the arrival of outcross pollen, can reduce annual seed set in predominantly bee-pollinated trees. However, various other factors may cause flower abortion in addition to pollen limitation. The results of this study would be very valuable to the seed orchard managers and the silviculturists to manage the seed production areas (SPAs) of G. arborea. The study ultimately recommends supplemental pollination in seed orchards of G. arborea for better genetic gain and good seed yield.     

Pollen flow,self-pollination and gender specialization: Factors affecting seed-set in the tristylous speciesLythrum salicaria (Lythraceae)

Some of the factors governing seed-set in three populations ofLythrum salicaria are examined during two seasons. Plants spatially segregated from legitimate neighbours by distances exceeding 1m set few seeds. Plants with small inflorescences set fewer seeds than those with medium or large inflorescences. Flowers having self pollen applied to the stigma before legitimate pollination set fewer seed per flower than those pollinated with legitimate pollen only. Seeds of the mid-style morph germinated more effectively than those of the long style morph and seeds of the short-style morph showed very low levels of germination.     

Plant size,geitonogamy and seed set in Ipomopsis aggregata

Summary We used powdered fluorescent dyes to estimate receipt of self vs. outcross pollen in the self-incompatible species Ipomopsis aggregata (Polemoniaceae). Flowers on small and large plants received equal amounts of outcross pollen, whereas flowers on large plants received more self pollen, so the proportion of self pollen delivered through geitonogamy increased with plant size. In natural populations emasculation of all flowers on a plant raised average seed set per flower from 5.19 to 6.99 and also raised fruit set, though not significantly. From these results one expects a negative correlation between plant size and seeds per flower. The opposite trend was observed in a sample of plants in the field, suggesting that deleterious effects of geitonogamy on female fecundity in large plants can be overruled by other factors such as size-related fruit or seed abortion. Results are discussed in relation to the evolution of gynodioecy.     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

Nectarless flowers with deep corolla tubes in Pedicularis: does long pistil length provide an arena for male competition?

Pollinator‐mediated selection does not seem to have a direct influence on the evolution of a long corolla tube in a nectarless flower. We hypothesized that the long pistil length of the nectarless flower with a deep corolla tube provided an opportunity for male competition. Pedicularis siphonantha, a nectarless and partially self‐incompatible lousewort with substantial variation in corolla tube length, was used to test the hypothesis. We tested whether and how corolla tube length affected seed production per capsule and seed germination rate with different pollination treatments. Flowers were hand‐pollinated with pollen from one self donor and one outcross donor and mixed pollen grains consisting of equal amounts from the two donor types, respectively. Additionally, seeds from open‐pollinated flowers with different corolla tube lengths were collected separately for measurement of germination rate. Pollination treatment and corolla tube length significantly affected number of seeds per capsule. Moreover, a significant positive relationship between seeds per capsule and corolla tube length was found when mixed hand pollination was conducted. Seeds of self hand‐pollinated flowers had a lower germination rate than those from outcross‐pollinated flowers. Under open pollination, seeds from flowers with longer corolla tubes tended to have higher germination rate. In P. siphonantha, outcross pollen may have a higher probability of contributing to the next generation when transferred to flowers with longer corolla tubes. The pistil length, therefore, should be seen as a female choice mechanism, which provides an arena for male‐to‐male competition. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 526–532.     

Bottlenecks and spatiotemporal variation in the sexual reproduction pathway of perennial meadow plants

Sexual reproduction is important for the growth of populations and the maintenance of genetic diversity. Several steps are involved in the sexual reproduction pathway of plants: the production of flowers, the production of seeds and the establishment of seedlings from seeds. In this paper we quantify the relative importance and spatiotemporal variability of these different steps for four grassland perennials: Centaurea jacea, Cirsium dissectum, Hypochaeris radicata and Succisa pratensis. We compared undisturbed meadows with meadows where the top soil layer had been removed as a restoration measure. Data on the number of flower heads per flowering rosette, the numbers of flowers and seeds per flower head, and the seedling establishment probabilities per seed were collected by field observations and experiments in several sites and years. Combination of these data shows that H. radicata and S. pratensis have higher recruitment rates (1.9 and 3.3 seedlings per year per flowering rosette, respectively) than the more clonal C. dissectum and C. jacea (0.027 and 0.23, respectively). Seedling establishment is the major bottleneck for successful sexual reproduction in all species. Large losses also occurred due to failing seed set in C. dissectum. Comparison of the coefficients of variation per step in space and time revealed that spatiotemporal variability was largest in seedling establishment, followed closely by flower head production and seed set.     

Reproductive conflicts ofPalicourea padifolia (Rubiaceae) a distylous shrub of a tropical cloud forest in Mexico

We studied a population of the distylousPalicourea padifolia (Rubiaceae) in a cloud forest remnant near Xalapa City, Veracruz, México to explore possible asymmetries between floral morphs in the attractiveness to pollinators, seed dispersers, nectar robbers, floral parasites, and herbivores. We first assessed heterostyly and reciprocal herkogamy by measuring floral attributes such as corolla length (buds and open flowers), style and anther heights, stigma and stamen lengths and the distance between the anther tip to the stigma lobe. We then estimated floral and fruit attributes such as flower size, anther height, number and size of pollen grains, fruit size, seed size, nectar production, and flower and fruit standing crops to assess differences between floral morphs in attracting and effectively using mutualistic pollinators and seed dispersers. Also, floral parasitism and nectar robbing were assessed in this study as a measure of flower attractiveness to antagonists. The system seems to conform well to classical heterostyly (e.g. reciprocal stamen/style lengths, pollen and anther dimorphism, intramorph incompatibility) yet, there were several tantalizing differences observed between pin and thrum morphs. Thrum flowers have longer corollas and larger but fewer pollen grains than pin flowers. Both morphs produced the same total number of inflorescences, developed the same number of buds, and opened the same number of flowers per inflorescence during the flowering season. Nectar production and sugar concentration were similar between floral morphs but the reward was not offered symmetrically to floral visitors throughout the day. Nectar concentration was higher in pin flowers in the afternoon. The numbers of developing, fully developed, and ripe fruits were the same between floral morphs, however, fruits and seeds were larger than those of thrums. The incidence of fly larvae was higher among thrum flowers and damage by nectar robbing was the same between floral morphs. Fruit abortion patterns of flowers manually pollinated suggest intra-morph sterility (self and intramorph incompatibility). There were no differences between morphs in fruit and seed set per flower following legitimate pollination although thrums were more leaky than the pins (intramorph compatibility).     

Floral display size in comfrey, Symphytum officinale L. (Boraginaceae): relationships with visitation by three bumblebee species and subsequent seed set

The fecundity of insect-pollinated plants may not be linearly related to the number of flowers produced, since floral display will influence pollinator foraging patterns. We may expect more visits to plants with more flowers, but do these large plants receive more or fewer visits per flower than small plants? Do all pollinator species respond in the same way? We would also expect foragers to move less between plants when the number of flowers per plant are large, which may reduce cross-pollination compared to plants with few flowers. We examine the relationships between numbers of inflorescence per plant, bumblebee foraging behaviour and seed set in comfrey, Symphytum officinale, a self-incompatible perennial herb. Bumblebee species differed in their response to the size of floral display. More individuals of Bombus pratorum and the nectar-robbing B.?terrestris were attracted to plants with larger floral displays, but B. pascuorum exhibited no increase in recruitment according to display size. Once attracted, all bee species visited more inflorescences per plant on plants with more inflorescences. Overall the visitation rate per inflorescence and seed set per flower was independent of the number of inflorescences per plant. Variation in seed set was not explained by the numbers of bumblebees attracted or by the number of inflorescences they visited for any bee species. However, the mean seed set per flower (1.18) was far below the maximum possible (4 per flower). We suggest that in this system seed set is not limited by pollination but by other factors, possibly nutritional resources.     

Sources of variation in pollinator contribution within a guild: the effects of plant and pollinator factors

Among plants visited by many pollinator species, the relative contribution of each pollinator to plant reproduction is determined by variation in both pollinator and plant traits. Here we evaluate how pollinator movement among plants, apparent pollen carryover, ovule number, resource limitation of seed set, and pollen output affect variation in contribution of individual pollinator species to seed set in Lithophragma parviflorum (Saxifragaceae), a species visited by a broad spectrum of visitors, including beeflies, bees and a moth species. A previous study demonstrated differences among visitor species in their single-visit pollination efficacy but did not evaluate how differences in visitation patterns and pollen carryover affect pollinator efficacy. Incorporation of differential visitation patterns and pollen carryover effects —commonly cited as potentially important in evaluating pollinator guilds — had minor effects (0–0.6% change) on the estimates of relative contribution based on visit frequency and single-visit efficacy alone. Beeflies visited significantly more flowers per inflorescence than the bees and the moth. Seed set remained virtually constant during the first three visited flowers for beeflies and larger bees, indicating that apparent pollen carryover did not reduce per-visit efficacy of these taxa. In contrast, Greya moth visits showed a decrease in seed set by 55.4% and the smaller bees by 45.4% from first to second flower. The larger carryover effects in smaller bees and Greya were diminished in importance by their small overall contribution to seed set. Three variable plant traits may affect seed set: ovule number, resource limitation on seed maturation, and pollen output. Ovule number per flower declined strongly with later position within inflorescences. Numbers were much higher in first-year greenhouse-grown plants than in field populations, and differences increased during 3 years of study. Mean pollen count by position varied 7-fold among flowers; it paralleled ovule number variation, resulting in a relatively stable pollen:ovule ratio. Resource limitation of seed set increased strongly with later flowering, with seed set in hand-pollinated flowers ranging from 66% in early flowers to 0% in the last two flowers of all plants. Variation in ovule number and resource limitation of seed maturation jointly had a strong effect on the number of seeds per flower. Visitation to early flowers had the potential to cause more seed set than visitation to later flowers. Overall, the most important sources of variation to seed production contribution were differences among pollinators in abundance and absolute efficacy (ovules fertilized on a single visit) and potentially differential phenology among visitor species. These effects are likely to vary among populations and years.     

Comparative fecundity of morphs in a mixed monoclinous-andromonoecious population ofWurmbea dioica subsp.alba (Liliaceae) in Western Australia

A population ofWurmbea dioica subsp.alba in Western Australia contained monoclinous and andromonoecious individuals in roughly equal proportions. The average number of flowers per inflorescence for the former was 2.6 and for the latter 2.9, with its terminal flower staminate. Ovule number and seed production per perfect flower of both morphs decreased progressively from lower to upper flowers in the inflorescence. Two-flowered monoclinous individuals had a greater percentage of ovules maturing to seed than did threeflowered ones, but seed production of the latter exceeded that of the former. No differences in percent seed set or in seed production were noted for similar individuals of andromonoecious individuals. Although monoclinous and andromonoecious plants apparently contributed equally to the pollen pool, seed production of the former exceeded that of the latter. Thus, the energetic costs of monocliny exceed those of andromonoecy in this population.