Takashi Y. Ida

Early onset of spring increases the phenological mismatch between plants and pollinators.

Climate warming accelerates the timing of flowering and insect pollinator emergence, especially in spring. If these phenological shifts progress independently between species, features of plant-pollinator mutualisms may be modified. However, evidence of phenological mismatch in pollination systems is limited. We investigated the phenologies of a spring ephemeral, Corydalis ambigua, and its pollinators (bumble bees), and seed-set success over 10-14 years in three populations. Although both flowering onset and first detection of overwintered queen bees in the C. ambigua populations were closely related to snowmelt time and/or spring temperature, flowering tended to be ahead of first pollinator detection when spring came early, resulting in lower seed production owing to low pollination service. Relationships between flowering onset time, phenological mismatch, and seed-set success strongly suggest that phenological mismatch is a major limiting factor for reproduction of spring ephemerals. This report demonstrates the mechanism of phenological mismatch and its ecological impact on plant-pollinator interactions based on long-term monitoring. Frequent occurrence of mismatch can decrease seed production and may affect the population dynamics of spring ephemerals.

Floral color change in Weigela middendorffiana (Caprifoliaceae): reduction of geitonogamous pollination by bumble bees

We examined the significance of retaining color-changed flowers in pollination success of Weigela middendorffiana through a single visit of bumble bees. Inner parts of flowers changed color with age from yellow to red. In an investigation of the mating system, duration of each color phase, reproductive ability of each of the color-phase flowers, and the effects of color-changed flowers on bumble bee behavior (1) flowers of this species were self-incompatible, (2) color-changed flowers provided little reward to pollinators and little residual reproductive ability, (3) the timing of floral color change was delayed with the progress of flowering season within individual plants, while the duration of the red phase shortened with the progress of flowering season, and (4) red-phase flowers did not attract bumble bees at a distance but did contribute to reducing the number of successive flower visits during a single stay within the plants. Red-phase flowers seemed to indicate the low reward level of old flowers and functioned as a cue to discourage pollinators from staying longer on the same plant. Our results predict that the retention of color-changed flowers without sexual function can enhance the pollination success of a whole plant through male function by reducing successive flower visits during a single stay of pollinators, i.e., geitonogamous pollination.

Photosynthetic compensation by the reproductive structures in the spring ephemeral Gagea lutea

Growth and reproduction of spring ephemerals inhabiting deciduous forests progress simultaneously during a short period from snowmelt to canopy closure. To clarify the mechanism to mitigate the cost of reproduction, contributions of foliar and non-foliar photosynthetic products to seed production were examined in a spring ephemeral Gagea lutea. Leaf growth, foliar and non-foliar photosynthetic activities, and total assimilated products were compared among reproductive-intact, floral bud-removal, and vegetative plants. Translocation of current photosynthetic products to individual organs was quantified by 13CO2-trace experiment. Bulb growth was compared between hand-pollination and floral bud-removal treatments. Finally, seed set was compared between intact, leaf-clipping, and bract-clipping treatments. Fruit-forming plants retained leaves longer than vegetative and floral bud-removal plants, but the assimilative contribution of extended leaf longevity was negligible. Carbon supply by bract photosynthesis was large enough for fruit development, while carbon supply by fruit photosynthesis was offset by the high respiration loss. Foliar photosynthetic products were largely transported to bulbs, while translocation to reproductive functions was negligible. Because the floral bud-removal increased the bulb growth, lack of reproduction could lead to more storage. The leaf-clipping had no effect on seed production, while the bract-clipping significantly reduced the seed production. Therefore, current photosynthesis of leafy bracts might be a major carbon source for fruit development. This self-compensative mechanism of reproductive structure enables the continuous reproductive activity in this species.

Demand‐driven resource investment in annual seed production by a perennial angiosperm precludes resource limitation

The limits on annual seed production have long been characterized as restriction by either pollination success or resource provision to seed development. This expected dichotomy between pollen and resource limitation is based on the assumption that reproductive resources are fixed, which is reasonable for semelparous species. In contrast, iteroparity can ease the constraints on reproductive output per breeding season, if resources can be either mobilized from past storage or borrowed against future performance. For perennial plants, these options allow enhanced reproductive investment in response to unusually good pollination, so that annual seed production may not be pollen or resource limited. We assessed demand-governed reproductive investment by manipulating both resource supply capacity (partial defoliation) and resource demand (pollination quality: fully self-pollination, fully cross-pollination, or combinations of partial self- and cross-pollination within the inflorescence) for a forest herb, Stenanthium occidentale, which is subject to strong pre-dispersal inbreeding depression. Insensitivity to partial defoliation indicated that reproductive output was not source regulated. Instead, demand by developing seeds governs resource distribution, as demonstrated by elevated photosynthate translocation to fruits on fully cross-pollinated plants and the ability of completely defoliated plants to produce seeds. Such contingent resource allocation eliminates a simple dichotomy between pollen receipt and resource availability as limits on annual seed production. Instead, such flexible reproductive investment allows iteroparous perennials to participate maximally in current reproduction (as determined by ovule production) following superior pollination, or to conserve resources for future reproduction following poor pollination.

Comparison of light harvesting and resource allocation strategies between two rhizomatous herbaceous species inhabiting deciduous forests

Light conditions on the floor of deciduous forests are determined by the leaf dynamics of canopy trees and gap formation. Such spatiotemporal variations of light availability should affect the resource partitioning strategies of understory herbs. Although rhizomatous species are common in understory, relationships between rhizome structure, vegetative growth, and sexual reproduction are unclear in terms of carbon allocation. We compared the photosynthetic characteristics and carbon translocation patterns in the under-canopy and light-gap sites between two summer-green perennial species: Cardamine leucantha with an annual long rhizome, and Smilacina japonica with a perennial short rhizome system. Flowering of both species occurs in early summer under decreasing light availability. In the light-gap, C. leucantha maintained high photosynthetic activity due to continuous leaf production, resulting in higher seed production than in the under-canopy. In contrast, the photosynthetic rate of S. japonica, producing leaves simultaneously, decreased with time irrespective of light conditions, resulting in stable seed production in both sites. Although seasonally decreasing light availability commonly restricts carbon assimilation of understory herbs, the responses of resource partitioning to variations in light availability depend greatly on the belowground structure of individual species.

Modification of bumblebee behavior by floral color change and implications for pollen transfer in Weigela middendorffiana.

Flowers of Weigela middendorffiana change the color from yellow to red. The previous study revealed that red-phase flowers no longer have sexual function and nectar, and bumblebees selectively visit yellow-phase flowers. The present study examined how retaining color-changed flowers can regulate the foraging behavior of bumblebees and pollen transport among flowers within (geitonogamous pollination) and between (outcrossing pollination) plants and how the behavior is influenced by display size (i.e., number of functional flowers) and visitation frequency. The visitation frequencies of bumblebees to plants and successive flower probes within plants were observed in the field using plants whose flower number and composition of the two color-phase flowers had been manipulated. To evaluate pollination efficiency over multiple pollinator visits, a pollen transport model was constructed based on the observed bumblebee behavior. In the simulation, three flowering patterns associated with display size and existence of color-changed flowers were postulated as follows: Type 1, large display (100 functional flowers) and no retention of color-changed flowers; Type 2, small display (50 functional flowers) and retention of color-changed flowers (50 old flowers), and; Type 3, large display (100 functional flowers) and retention of color-changed flowers (100 old flowers). Color-changed flowers did not contribute to increasing bumblebee attraction at a distance but reduced the number of successive flower probes within plants. Comparisons of pollen transfer between Types 1 and 3 revealed that the retention of color-changed flowers did not influence the total amount of pollen exported when pollinator visits were abundant (>100 visits) but decreased geitonogamous pollination. Comparisons between Types 2 and 3 revealed that the discouragement effect of floral color change on successive probes accelerated in plants with a large display size. Overall, the floral color change strategy contributed to reduce geitonogamous pollination, but its effectiveness was highly sensitive to display size and pollinator frequency.

Effects of defoliation and shading on the physiological cost of reproduction in silky locoweed Oxytropis sericea

BACKGROUND The production of flowers, fruits and seeds demands considerable energy and nutrients, which can limit the allocation of these resources to other plant functions and, thereby, influence survival and future reproduction. The magnitude of the physiological costs of reproduction depends on both the factors limiting seed production (pollen, ovules or resources) and the capacity of plants to compensate for high resource demand. METHODS To assess the magnitude and consequences of reproductive costs, we used shading and defoliation to reduce photosynthate production by fully pollinated plants of a perennial legume, Oxytropis sericea (Fabaceae), and examined the resulting impact on photosynthate allocation, and nectar, fruit and seed production. KEY RESULTS Although these leaf manipulations reduced photosynthesis and nectar production, they did not alter photosynthate allocation, as revealed by (13)C tracing, or fruit or seed production. That photosynthate allocation to reproductive organs increased >190 % and taproot mass declined by 29 % between flowering and fruiting indicates that reproduction was physiologically costly. CONCLUSIONS The insensitivity of fruit and seed production to leaf manipulation is consistent with either compensatory mobilization of stored resources or ovule limitation. Seed production differed considerably between the two years of the study in association with contrasting precipitation prior to flowering, perhaps reflecting contrasting limits on reproductive performance.

The consequences of demand-driven seed provisioning for sexual differences in reproductive investment in Thalictrum occidentale (Ranunculaceae)

Summary Many iteroparous angiosperms may benefit from flexible annual resource allocation in response to variable reproductive opportunities induced by external conditions. If maximal reproductive investment is fixed, lack of reproductive sinks would cause resource redistribution to other sinks. Alternatively, reproductive investment may vary depending on the demand of reproductive sinks, changing source–sink relations. In particular, differential responses by males and females to the demands of flower and seed production may cause sexual dimorphism. We assess the occurrence of demand-driven seed allocation by females and its implications for sexual differences in reproductive investment, including the dynamics of mass and carbon allocation and the physiological cost of reproduction, for a dioecious, perennial herb, Thalictrum occidentale. We specifically quantified allocation responses to partial defoliation, which reduced current resource supply, and partial flower/fruit removal, which reduced the aggregate demand of reproductive sinks. During flowering, males preferentially invested carbon and mass in flowers at the expense of vegetative organs, whereas females allocated less mass to flowers and invested more in new rhizome production for future performance than males. In contrast, during early fruiting, both sexes had new rhizomes of similar size and a doubling of reproductive mass by females after flowering resulted in similar total reproductive investment for both sexes. Manipulation of the source–sink balance did not influence carbon allocation, except that partial fruit removal increased new rhizome mass compared with intact plants. Females with many fertilized ovules invested proportionally more in seed number and mass per seed than females with few fertilized ovules, indicating both demand-driven seed maturation and its elimination of seed size-number trade-off. Furthermore, males consistently exhibited size-dependent flower production, whereas females exhibited size-dependent flower production only if they had not reproduced during the previous year. Synthesis. This study demonstrates that sexual differences in policies of reproductive investment and the timing of the physiological costs of reproduction impose contrasting allocation schedules. Males invested in reproduction proportionally with their size, whereas females invested flexibly in seeds in response to the demand of developing embryos. Thus, the contrasting certainty and timing of reproductive resource requirements between the sexes contribute to sexual dimorphism.