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Abstract: Clonal and cutting stem morphology variations in rooting ability were simultaneously examined in Allanblackia floribunda Oliv. using leafy stem cuttings in two experiments. The hypothesis tested is that clonal [...]

Author(s): Wudan Yan [1] Nathan Lakey practically has to shout to be heard over the whirring machinery in the laboratory at Orion Biosains near Kuala Lumpur. One source of the [...]

Abstract: The increasing use of exotic and hybrid poplar species in forestry and the lack of genetic barriers between most poplar species may present a risk to the genetic integrity [...]

A tradeoff between growth and reproduction, often inferred from an inverse correlation between these two variables, is a fundamental paradigm of life-history evolution. Oak species provide a unique test of this relationship because different species mature acorns either in the year of pollination or in the year after pollination. This difference allows for an interspecific comparison testing whether the apparent tradeoff is causal or the result of confounding factors influencing growth and reproduction independently. Based on 13 years of data on five California oak species, we found significant negative correlations between radial growth and seed production in the three species that produce acorns the same year in which pollination occurs, but not in two species that mature acorns the year after pollination. Rainfall, which correlates positively with radial growth and correlates negatively with acorn production (based on the year of pollination), appears to be driving this pattern. We conclude that the observed negative correlations are not causal, but rather a consequence of growth and reproduction being dependent, in opposite ways, on environmental conditions. Thus, contrary to the current consensus, growth and reproduction in these species are apparently largely independent of each other. In contrast, tradeoffs between current and future reproduction appear to be much more important in the life-history evolution of these long-lived plants. We also conclude that a negative correlation does not necessarily imply a causal mechanism and should not be used as the only evidence supporting a tradeoff. allocation | cost of reproduction | life-history evolution | reproductive effort | masting

This study examines how the latitude of cultivation of Ginkgo biloba affects the timing of all phases of its sexual reproductive cycle, from pollination through germination. Seeds produced by trees growing in warm-temperate climates germinate earlier in the year than seeds produced in cold-temperate climates, and they have a longer period of time available for seedling establishment. The embryos of G. biloba seeds possess a temperature-dependent developmental-delay mechanism that allows seeds to survive winter by preventing premature germination in the fall. This and other cold-climate adaptations appear to have evolved within the genus Ginkgo during the early Cretaceous, when the Northern Hemisphere was undergoing dramatic cooling after a long period of stable, warm conditions. Ginkgo biloba seeds possess an odoriferous sarcotesta that attracts mammalian scavengers in Asia--most notably members of the Carnivora--presumably by mimicking the smell of carrion. Seeds cleaned of their sarcotesta germinated faster and at higher percentages than those with their sarcotesta intact, suggesting that animal dispersal plays an important role in promoting seedling establishment. During the Cretaceous, potential dispersal agents included mammals, birds, and carnivorous dinosaurs., Abstract Introduction Literature Review Pollination and Fertilization Germination Materials and Methods Results Discussion Evolutionary Implications Ginkgo Seed Dispersal Acknowledgments Literature Cited Introduction Ginkgo biloba L. is a large tree native [...]

Plant somatic cells have the remarkable ability to regenerate an entire organism. Many species in the genus Kalanchoe, known as 'mother of thousands,' develop plantlets on the leaf margins. Using key regulators of organogenesis (STM) and embryogenesis (LEC1 and FUS3) processes, we analyzed asexual reproduction in Kalanchoe leaves. Suppression of STM abolished the ability to make plantlets. Here, we report that constitutive plantlet-forming species, like Kalanchoe daigremontiana, form plantlets by coopting both organogenesis and embryogenesis programs into leaves. These species have a defective LEC1 gene and produce nonviable seed, whereas species that produce plantlets only upon stress induction have an intact LEC1 gene and produce viable seed. The latter species are basal in the genus, suggesting that induced-plantlet formation and seed viability are ancestral traits. We provide evidence that asexual reproduction likely initiated as a process of organogenesis and then recruited an embryogenesis program into the leaves in response to loss of sexual reproduction within this genus. embryogenesis | LEC1 | STM

Floral sex allocation at the individual and first-order branch levels and the relation between these levels were examined in Betula platyphylla var. japonica, a wind-pollinated monoecious tree. Floral sex allocation at the individual level varied with resource availability in a pattern similar to that predicted by the Masaka and Takada model (Journal of Theoretical Biology 240: 114-125). Thus, individual trees with few reproductive resources produced only female or male inflorescences, whereas individuals with many resources rarely had a high male ratio (i.e., number of male inflorescences/total number of inflorescences). Furthermore, the number of male inflorescences tended to reach an upper limit, whereas the number of female inflorescences increased monotonically with increasing reproductive investment. The patterns of floral sex allocation at the first- order branch level were analogous to those at the individual level. Thus, each first-order branch of B. platyphylla var. japonica behaves like an individual, and the floral sex allocation of a given branch does not necessarily represent the individual tree. The effect of the individual-level floral sex ratio on branch-level floral sex allocation indicates that branch behavior is controlled by the individual. Key words: first-order branch; floral sex allocation: individual; reproductive resources; wind-pollinated monoecy.

CERES-Maize simulates kernel set as a sourcelimited process based on average plant growth rate during the lag phase after flowering. Yet the number of kernels formed by maize (Zea mays L.) also depends on timely interaction between male and female flowers, which can limit formation of reproductive sinks under some conditions. Failure to account for sink-limited kernel set may contribute to simulation error observed under conditions that affect dynamics of pollen shed or silking, but do not alter crop growth rate. We developed algorithms for a Flowering Model to simulate sink-limited kernel set from flowering dynamics. This model was calibrated against kernel production in hybrid seed production fields and then linked to CERES-Maize. The Modified CERES-Maize was calibrated against two years of field data involving three hybrids, eight population densities, and seven N levels. Integrating the capacity to simulate sinklimited kernel set with source-limited kernel set increased simulation accuracy dramatically, relative to original CERES-Maize. For 13 commercial fields tested, Modified CERES-Maize decreased simulation error for kernels per plant from 17.1 to 2.3%, improved P between measured and simulated values from 0.77 to 0.87, and decreased simulation error indicators mean error, root mean square error, and mean square deviation by 85, 40, and 64%, respectively. Modified CERES-Maize accounts for a much greater range of variability in the biological processes controlling kernel set.

Invasive species are a global threat to biodiversity and the functioning of natural ecosystems. Here, we report on a two-year experiment aimed at elucidating the combined and relative effects of three key controls on plant invasions: propagule supply, soil nitrogen (N) availability, and herbivory by native insects. We focus on the exotic species Lespedeza cuneata, a Rank 1 invasive species. Propagule supply and soil N-availability interacted to control the density and foliar cover of L. cuneata. In low N plots, density and foliar cover of L. cuneata were higher in the propagule addition plots than in the plots to which propagules were not added. Surprisingly, this interaction was significant only when the abundance of herbivores was experimentally reduced. This experiment provides evidence that native insect herbivores mediate the interactive effects of propagule supply and resources on invasion by a widespread invasive plant species. Key words: biological invasion; dispersal; herbivory; invasive species; Lespedeza cuneata; nutrients: plant-insect interactions.

We conducted a cytogenetic study of sexual lines of Boechera stricta and Boechera holboellii (2n = 14) and seven diploid apomictic accessions of their interspecific hybrid Boechera divaricarpa and B. holboellii (2n = 14 or 15). By studying chromosome morphology, rDNA repeats, genome painting, male meiosis, pollen morphology, and flow-cytometry seed screens, we revealed an unexpected plethora of chromosome forms, pairing behavior, and hybrid composition in all apomictic lines. Genome painting demonstrated that the apomicts are alloploid with variable numbers of B. stricta and B. holboellii-like chromosomes. We assume that large-scale homeologous chromosome substitutions took place in the apomictic hybrids that resulted from recurrent diploid-polyploid transitions through restitutional meiosis and polyploidy-diploid transitions through reductional meiosis. A second peculiarity was the presence of a largely heterochromatic chromosome (Het) in all apomictic accessions (2n = 14 and 15) and an additional smaller chromosome (De/) in the aneuploids (2n = 15). Both chromosomes share repetitive pericentromere repeats with those from the sexual B. stricta, suggesting that they originated from this species. Pairing and behavior at meiosis I of the Het share features with both Y and B chromosomes and suggest that the Del arose from a translocation event or homeologous recombination between a B. holboellii (or related taxon) and a B. stricta chromosome. Based on its presence exclusively in apomictic accessions, we propose that the Het chromosome plays a role in the genetic control of apomixis. apomixis | chromosome evolution | genome painting | heterochromatic chromosome | homeologous substitution

To increase our knowledge about mating-system evolution, we need to understand the relationship between specific floral traits and mating system. Species of Collinsia (Plantaginaceae) vary extensively in mating system; this variation is associated with variation in floral morphology and development and with the timing of self-pollination. Counterintuitively, large-flowered, more outcrossing species tend to have delayed stigma receptivity, reducing the amount of time that the stigma is receptive to cross-pollination before autonomous self-pollination. To understand how the timing of stigma receptivity is related to mating-system evolution, we studied in detail the timing of both stigma receptivity and self-pollination (anther-stigma contact) in two greenhouse-grown populations of large-flowered Collinsia heterophylla. Crosses on emasculated flowers at different stages of floral development always produced seeds, suggesting that cross-fertilization can be effected by pollen arriving prior to physiological receptivity. Phenotypic and genetic variation within populations in the timing of stigma receptivity and anther-stigma contact was substantial, although slightly less for the contact. Despite strong interspecific and interpopulation correlations, we did not find an among-genet phenotypic correlation between the traits. This indicates that each trait may respond independently to selection, and the trait association may be the result of correlational selection. Key words: Collinsia heterophylla; delayed selfing; delayed stigma receptivity; evolvability: flower development; beritability; mixed mating system; Plantaginaceae.

Abstract: Outcrossing, wind-pollinated members of the Pinaceae have high self-pollination rates yet produce few selfed seedlings. How are selfs selectively eliminated? Barriers to selfing have long been considered to be [...]

Bahiagrass (Paspalum notatum Flugge) is one of the major forage grasses used for beef cattle production in the southern USA. Description of the reproductive characteristics of bahiagrass germplasm is needed for genetic improvement of this forage crop. The objective was to determine and compare the mode of reproduction of diploid and tetraploid germplasm of bahiagrass. Reproductive behavior was determined by embryo sac observations and a series of controlled pollination studies in the greenhouse and field in 2004 and 2005. Two sexual diploid populations ('Pensacola' and 'Tifton 9') were not different in self- or cross-fertility, indicating that the general fertility of the crop was not affected by phenotypic mass selection. Sexual diploids, as a group, were determined to be primarily cross-pollinated, with low but variable levels of self-fertility. A group of 20 artificially induced tetraploids was determined to be primarily sexual and cross-pollinated. Fertility of these autotetraploids was lower than the original diploids. Two selected sexual tetraploids produced similar amounts of seed when self- or cross-pollinated. In contrast, six other tetraploids, including the cultivars Argentine and Wilmington, the experimental hybrid Tifton 7, and three plant introductions 315732, 315733, and 315734 reproduced primarily by apomixis. The sexual or apomictic expression, the general fertility, and other agronomic characteristics of these groups of tetraploids define their usefulness for genetic improvement research.

Duration of the reproductive phase (DRP) is critical for soybean [Glycine max (L.) Merr.] yield. Manipulation of this phase may benefit breeding for higher yield. The soybean E-gene series control time to flowering and maturity through a photoperiod-mediated response. It is possible that E-genes and photoperiod may control the DRP. Two 2-yr studies were conducted with the objective to assess whether the DRP is influenced by either or both E-gene alleles and post-flowering photoperiod. In a planting-date experiment, the main plots were two planting dates, and sub plots were 15 E-gene near-isogenic lines (NILs) in two genetic backgrounds. In a daylength-extension experiment, seven E-gene NILs were planted to synchronize flowering date. Two photoperiods treatments were imposed after flowering: (i) ambient and (ii) ambient plus 3-h incandescent daylength extension. Early planting and daylength extension exposed plants to a longer post-flowering photoperiod, and resulted in an increase in the DRP. When exposed to the same post-flowering photoperiod, the DRP was positively related to the number of dominant E-gene alleles. Both E-genes and post-flowering photoperiod controlled the DRP and this was in addition to the known impact of E-genes on time to flowering. Hence, E-genes control time to maturity through both their effect on time to flowering and through their effect on the DRP. The results of this study underline the importance of time of flowering for determination of the DRP.

Abstract: A total of 2561 slash pine (Pinus elliottii Engelm. var. elliottii) grafts were topgrafted in the winter of 2003 and evaluated in January of 2004. The objectives of this [...]

Abstract: Unlike their diploid relatives, some triploid and tetraploid Crataegus frequently produce unreduced megagametophytes. In all cases, pollination is required for successful seed set, but in polyploids, endosperm formation can [...]

Synchronous pollination improves maize (Zea mays L.) kernel set, but the physiological reasons behind this response are yet unclear. We registered ovary fresh weight evolution at three floret positions along the ear (base, middle and tip) of plants subjected to natural (NP) or synchronous (SP) pollination during two growing seasons. Synchronous pollination of ears bagged before silking was performed five days after silking (DAS). Ovary weight increased exponentially at all floret positions (P [less than or equal to] 0.01), but a lag in this trait was detected among those at the base and middle of SP ears. At each sampling date, florets along the ear differed (P < 0.01) in ovary weight, but differences were larger for NP than for SP plants. At pollination of each floret position, however, the range in ovary weight was smaller in NP ears (ca. 4 to 5 mg [ovary.sup.-1]) than in SP ears (4 to 9 mg [ovary.sup.-1]). Contrarily, just pollinated ovaries at the tip of the ear of NP plants (i.e. on ca. 4-5 DAS) experienced the high growth rate of those already pollinated at the base (ca. 0.7 mg [d.sup.-1] for the former and ca. 3.7 mg [d.sup.-1] for the latter). The range in ovary growth rate along the ear was drastically reduced under synchronous pollination on 5 DAS (ca. 0.55 mg d 1 for the base and ca. 0.47 mg [d.sup.-1] for the tip). The larger the tip-to-base ratio in ovary growth rate, the larger the number of kernels set per plant ([r.sup.2] = 0.94; P = 0.03).

The relationship between net primary productivity and biological diversity has been a central topic in ecology for several decades. The unimodal ('hump-back') relationship has been the most widely accepted for plants with the decrease in diversity at high productivity usually attributed to competitive exclusion. However, the relatively small species pool size under high productivity conditions may account for this pattern as well. Small species pool sizes for highly productive habitats are characteristic of temperate regions, where productive habitats for speciation and species migration have historically been rare. In contrast, productive habitats in the tropics have been relatively common during evolutionary history, resulting in large species pools. We hypothesize that evolutionary history contributes to the observed productivity--diversity relationship of plants, and that the productivity--diversity relationship differs between temperate and tropical regions. We investigated the productivity--diversity relationship patterns from 163 case studies throughout the world. Latitude described ~80% of the variation in the shape of the relationships. The unimodal relationship was found to dominate in the temperate zone, whereas the positive relationship was significantly more common in the tropics. We detected no influence due to methods of productivity measurement, but unimodal or positive productivity--diversity relationships were more likely within larger ranges of productivity. The length of the productivity gradient did not affect the latitudinal influence. In summary, the shape of the productivity--diversity relationship differs between temperate and tropical regions and the different evolutionary history of the local species pools is a probable cause for the difference. Key words: biodiversity conservation; generality of ecological rules; global diversity patterns; latitudinal gradient; macroecology; meta-study; plant biomass; plant species richness; primary productivity; productivity--diversity relationship; species pool; tropical ecology.

Plant fertility is a central subject of many questions in plant evolutionary and conservation biology. Pollen availability, abiotic resources, and flowering pattern can limit fruit and seed production. Open pollination and pollen supplementation studies are used to estimate any pollen limitation in natural populations. To study the impact of these factors on the reproductive success of Vriesea gigantea, an epiphytic bromeliad in the Atlantic Rainforest in Brazil, its fertility in four natural populations in Itapua State Park was assessed by considering plant and inflorescence size, flower production, fruit and seed set, flower and fruit set pattern, and seed viability and germination rate. Supplemental pollination in adult plants was used to determine whether fruit production in V. gigantea is limited by reception of pollen. The results showed that V. gigantea has a high production of flowers, fruits, and seeds. Seeds are highly viable in all populations, presenting an average germination rate of 94% (SE [+ or -] 3.5). Plants of V. gigantea from Itapua State Park are highly fertile. The high proportion of fruit and seed set after manual hand pollination indicates that the species is self-compatible. Pollination treatments showed evidence of pollinator limitation in the Itapua State Park population. Key words: Bromeliaceae; flower production; fruit set; Itapua State Park; pollen limitation; seed viability; self-compatibility.

Nectar production may disproportionately benefit male relative to female pollination success. In such cases, sexual selection is often suggested as the cause of asymmetric benefits, yet sexual selection in plants--particularly plants with hermaphroditic flowers--is infrequently tested empirically. Here, I used a protandrous herb with male-biased nectar production (Chrysothemis friedrichsthaliana, Gesneriaceae) to test predictions from sexual selection theory. During three flowering seasons, I measured nectar production, pollinator visits, and male and female fecundity following different numbers of cross-pollination events. In accordance with sexual selection predictions, (1) nectar production was greater during the male phase by at least 65%; (2) visits by the main pollinator (hummingbird Phaethornis striigularis) were limiting for part of the season, indicating that plants had to compete for pollinator visits; (3) pollinators spent 53% more time per visit and made 86% more visits to male- vs. female-phase flowers, suggesting that nectar increased male more than female pollination success; and (4) female fecundity was maximized by one visit, whereas male fecundity continued to increase with additional visits. Autonomous self-pollination further reduced visit requirements for maximum female seed set. These findings match specific sexual selection predictions: they link an observable male bias in a secondary sexual trait (nectar) to positive responses of mating participants (pollinators), resulting in more mating opportunities for mate-limited males, relative to apparently resource-limited females. This field-testing of theoretical predictions provides unique evidence that sexual selection helps maintain nectar production patterns in this and, quite likely, other hermaphroditic plant species. Key words: Chrysothemis friedrichsthaliana; Costa Rica; Gesneriaceae; hummingbird pollination; nectar; Phaethornis striigularis; pollinator-mediated selection; sexual selection.

Abstract: Pollen-mediated gene flow was analyzed in a small, fragmented, natural population of Fagus crenata Blume by surveying five microsatellite markers in seedlings derived from open-pollinated crosses. Paternity of 162 [...]

Abstract: Floral longevity is an important feature of a plant's reproductive strategy. The goal of this study was to examine flower life span in the shrub flame azalea (Rhododendron calendulaceum [...]

The article suggests a new theory for the evolution of overproduction of ovules in flowers. By overproducing ovules, females select fertilized ovules having similar resource absorption ability to produce seeds of uniform size.

Natural hybridization plays a critical role in speciation, the maintenance of reproductive isolation, and genetic introgression. While many plant species have hybrid swarms in areas of sympatry, the lack of hybrids among closely related sympatrically distributed species suggests that strong pre- and/or postzygotic barriers exist to hybridization. Gelsemium sempervirens and G. rankinii (Gelsemiaceae) are sympatrically distributed southeastern sister taxa that have strong postzygotic barriers to hybrid formation and high levels of genetic differentiation. In this study, two sympatric populations in Lowndes County, Georgia were surveyed from 1999-2005 to assess the role of temporal and pollinator isolation as potential prezygotic barriers. The populations had mostly non-overlapping flowering periods in 2003-2005, with significant differences in time of peak flowering and length of flowering. Both species shared a similar community of flower visitors, with the apid bee Habropoda laboriosa the dominant visitor to both species. A choice experiment found that H. laboriosa visited both species but preferred G. sempervirens. The primary prezygotic barrier is temporal isolation preventing hybridization in spite of the shared pollinators. This study suggests that reliance on a shared pollinator during speciation may limit opportunity for divergent selection on flowering time. Key words: floral isolation; Gelsemiaceae; Gelsemium; hybridization; prezygotic reproductive isolation; sympatry; temporal isolation.

When more pollen is present on stigmas than needed to fertilize all ovules, selection among pollen grains may occur due to effects of both pollen donors and maternal plants. We asked whether increasing plant age and flower age, two changes in maternal condition, altered the pattern of seed paternity after mixed pollination. We also asked whether changes in seed paternity affected offspring success in an experimental garden. While flower age did not affect seed paternity, there was a dramatic shift in pollen donor performance as plants aged. These differences were seen in the offspring as well, where the offspring of one pollen donor, which sired more seeds on young plants, flowered earlier in the season, and the offspring of another pollen donor, which sired more seeds on old plants, flowered later in tile season. Thus, change in maternal condition resulted in altered seed paternity, perhaps because the environment for pollen tube growth was different. The pattern of seed paternity and offspring performance suggests that pollen donors may show temporal specialization. Key words: Brassicaceae; flower age; mate choice; offspring fitness; plant age; pollen competition; sexual selection; wild radish.

Central to the ecology and evolution of a broad range of plants is understanding why they routinely have submaximal reproduction manifested as low seed: ovule and fruit : flower ratios. We know much less about the processes responsible for low seed : ovule ratios than we do for fruit : flower ratios. Current hypotheses for low seed : ovule ratios are largely drawn from those for fruit : flower ratios, including proximate (ecological) causes of pollen limitation, resource limitation, and pollen quality, as well as the ultimate (evolutionary) hypothesis of 'bet hedging' on stochastic pollination. Yet, such mechanisms operating on fruit : flower ratios at the whole-plant level may not best explain low seed : ovule ratios at the individual-flower level. We tested each of these proximate and ultimate causes for low seed : ovule ratios using the specialized pollination mutualism between senita cacti (Pachycereus schottii) and senita moths (Upiga virescens). Seed : ovule ratios were consistently low (~0.61). Such excess ovule production by senita likely has a strong genetic component given the significant differences among plants in ovule number and the consistency in ovule production by plants within and among flowering seasons. Excess ovule production and low seed : ovule ratios could not be explained by pollen limitation, resource limitation, pollen quality, or bet hedging. Nevertheless, phenotypic selection analyses did show significant selection gradients for increased ovule number, suggesting that other evolutionary processes may be responsible for excess ovule production and low seed : ovule ratios. In contrast, low fruit : flower ratios at the whole-plant level were explained by an apparent equilibrium between pollen and resource limitation. Thus, mechanisms responsible for low fruit : flower ratios at the whole-plant level are not necessarily in accord with those of low seed : ovule ratios at the individual-flower level. This suggests that we may need to adopt a more pluralistic approach to seed : ovule ratios and consider alternative hypotheses, including a greater array of proximate and ultimate causes. Initial results of this study suggest that floral allometry, selection on correlated floral traits, stigma clogging with pollen grains, and style clogging with pollen tubes may provide promising avenues for understanding low seed : ovule ratios. Key words: bet hedging; Cactaceae; floral allometry; fruit : flower ratio; Pachycerus schottii; phenotypic selection; pollen limitation; pollen load; pollen quality; resource limitation; seed : ovule ratio; senita.

In facultatively sexual species, lineages that reproduce asexually for a period of time can accumulate mutations that reduce their ability to undergo sexual reproduction when sex is favorable. We propagated Saccharomyces cerevisiae asexually for ~800 generations, after which we measured the change in sexual fitness, measured as the proportion of asci observed in sporulation medium. The sporulation rate in cultures propagated asexually at small population size declined by 8%, on average, over this time period, indicating that the majority of mutations that affect sporulation rate are deleterious. Interestingly, the sporulation rate in cultures propagated asexually at large population size improved by 11%, on average, indicating that selection on asexual function effectively eliminated most of the mutations deleterious to sporulation ability. These results suggest that pleiotropy between mutations' effects on asexual fitness and sexual fitness was predominantly positive, at least for the mutations accumulated in this experimental evolution study. A positive correlation between growth rate and sporulation rate among lines also provided evidence for positive pleiotropy. These results demonstrate that, at least under certain circumstances, selection acting on asexual fitness can help to maintain sexual function.

Abstract: Mating system pattern (selfing or outcrossing and correlated matings levels) comparisons between two interior spruce seed orchard designs (clonal-row and random) managed under intensive crown and pollen management were [...]

Pollination commonly limits seed production, as addition of pollen to stigmas often increases fecundity. This response is usually interpreted as evidence that plants' stigmas receive too few pollen grains to maximize ovule fertilization (quantity limitation); however, many genetic studies demonstrate that poor-quality pollen can also reduce seed production (quality limitation). We explore both aspects of pollen limitation theoretically with a dose-response model that incorporates a saturating negative-exponential relation of seed production to pollen receipt. This relation depends on aspects of ovule production, pollen import, pollen-pistil interactions and seed development, all of which can contribute to pollen limitation. Our model reveals that quantity limitation is restricted to the lowest range of pollen receipt, for which siring success per pollen grain is high, whereas quality limitation acts throughout the range of pollen receipt if plants do not import the highest-quality pollen. In addition to pollinator availability and efficiency, quantity limitation is governed by all post-pollination aspects of seed production. In contrast, quality limitation depends on the difference in survival of embryos sired by naturally delivered pollen vs. by pollen of maximal quality. We briefly illustrate the distinction between these two components of pollen limitation with results from the mistletoe Tristerix corymbosus. Our model also shows that the standard pollen-supplementation technique neither estimates the total intensity of pollen limitation nor distinguishes between its quantity and quality components. As an alternative, we propose a methodological protocol that requires both measurement of seed production following excess pollination with only outcross pollen and quantification of the dose-response relation of seed output to pollen receipt. This method estimates both the total extent of pollen limitation and its two components. Finally, we consider the influences on quantity and quality limitation, which reveals that quantity limitation probably occurs much less often than has been inferred from pollen-supplementation experiments. These interpretations suggest that an expanded perspective that recognizes the fecundity consequences of pollination with poor-quality pollen would promote ecological understanding of pollen limitation. Key words: plant reproduction; pollen limitation; pollen quality; pollination; seed production; supplemental pollination; Tristerix corymbosus.