Is analysing the nitrogen use at the plant canopy level a matter of choosing the right optimization criterion?

Herbivores can have strong deleterious effects on plant growth, reproduction, and even survival. Because these effects might be strongly interrelated, the direct consumptive effects of herbivores and a variety of indirect effects are difficult to untangle. Reductions in growth, for example, may strongly impact the flowering behaviour of plant species in the current season, but at the same time incur costs to survival, growth and reproduction in the next growing season(s). To get better insights in the effects of herbivory on the flowering behaviour of the long-lived polycarpic grassland herb Primula veris L., flowering patterns were monitored over ten consecutive years under two treatments (grazing and control mowing regimes). We tested the hypothesis that the size at flowering was affected by the presence of herbivores, and whether this translated into costs to future reproduction and survival. Overall, grazed plants were significantly smaller than control plants, and the size at which plants flowered was also significantly smaller when herbivores were present. The transition probability of flowering and of surviving into the next year was significantly smaller for all plants in the current year if they had been grazed than if they had been mown, indicating that herbivory incurred costs to both flowering and survival. Grazed plants also needed longer to start flowering, had fewer flowers and flowered less frequently, causing a significantly lower proportion of flowering adults in the population. These results suggest that the observed regression in plant size due to herbivory does not allow plants to capture enough resources to guarantee regular flowering in the longer run.     

Effects of supplementary feeding on growth and reproduction of three carnivorous plant species in a subarctic environment

1 The effect of supplementary feeding on growth and reproduction of three carnivorous plants species was investigated over a 6‐year period. Pinguicula alpina , P. villosa and P. vulgaris populations growing at two altitudes in a subarctic environment were fed with fruit flies ( Drosophila melanogaster ).
2 Fed plants increased in size relative to control plants during the first years. Subsequently a stable size difference between feeding levels was established. The weight of the over‐wintering part was higher in fed plants than in control plants.
3 The flowering frequency (i.e. proportion of plants in a population carrying flowers) was also higher in the fed plants. The proportion of flowering plants increased in the feeding treatment compared to control plants during more‐or‐less the whole experimental feeding period (5–6 years). Seed production also increased slowly in response to feeding.
4 No feeding effect on nitrogen and phosphorus concentrations in seeds and over‐wintering parts was found.
5 In most characteristics the high‐altitude populations were less responsive than populations growing at low altitude. In the high‐altitude population reproduction failed repeatedly, suggesting that seed output may be more dependent on abiotic factors than on resource availability in cold environments.
6 Pinguicula villosa and P. vulgaris used the new resources the same summer for increased leaf rosette growth and current reproduction, showing these species to be income breeders, while the third species ( P. alpina ) allocated a larger proportion to storage and future reproduction, characteristic of a capital breeder.     

Climate, size and flowering history determine flowering pattern of an orchid

The flowering pattern of plant species, including orchid species, may fluctuate irregularly. Several explanations are given in the literature to explain that pattern, including: costs associated with reproduction, herbivory effects, intrinsically triggered unpredictable variation of the system, and external conditions (i.e. weather). The influence of age is discussed, but is difficult to determine because relevant long-term field observations are generally absent in the literature. The influence of age, size, reproductive effort and climatic conditions on flowering variability of Himantoglossum hircinum are examined using data collected in a long-term project (1976–2001) in Germany. PCA and multiple regression analysis were used to analyse variability in flowering pattern over the years as a function of size and weather variability. We studied future size after flowering to quantify costs of reproduction. Flowering probability was strongly determined by plant size, while there was no significant influence of age class on flowering probability of the population. Costs associated with reproduction resulted in a decrease in plant size, causing reduced flowering probability of the plants in the following year. The weather explained about 50% of the yearly variation in the proportion of large plants and thus had an indirect, strong influence on the flowering percentage. We conclude that variability in flowering is caused mainly by the variability of weather conditions in the previous and current year, whereby reproductive effort causes further variability in flowering at the individual and, consequently, the population levels.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 511–526.     

Individual variation and the effects of weather, age and flowering history on survival and flowering of the long-lived perennial Gentiana pneumonanthe

Gentiana pneumonanthe , the marsh gentian, is a declining species in both Britain and Europe as a result of loss of its heathland habitat or inappropriate management of that which remains. We analysed long-term demographic data sets (1977-1991) from four populations of the plant to test the hypotheses that an individual plant's survival and its chances of flowering in any year are related to its age and performance in previous years, taking into account the climatic conditions that existed in those years The results show that: 1)'new'plants (2 yr old) had a higher mortality rate (mean = 20.8%) than'young'(3 yr old) plants (mean = 5.0%), which in turn had a higher mortality than older, 'mature' (four or more year old) plants (mean = 3.8%): 2)'new' plants had a higher mortality rate after either a growing season with below average rainfall or an above average winter rainfall; 3) mortality of'mature'plants was independent of age, indicating no evidence of senescence; 4)'mature'plants had a higher mortality rate after above average rainfall during the previous winter; 5) plants that have flowered within the previous two years are most likely to flower in the current year indicating high individual variation in flowering performance- 6) more plants flower and flowering plants produced more flowers in the two years following a warmer than average growing season.
The above factors are related to current site management and the implications are discussed. Suggestions are given for changes in management of the sites where this rare plant occurs.     

Tolerance of a perennial herb, <Emphasis Type="Italic">Pimpinella saxifraga</Emphasis>, to simulated flower herbivory and grazing: immediate repair of injury or postponed reproduction?

Perennial, polycarpic herbs can respond to herbivory either by (1) regrowth in the same season in order to compensate for lost reproductive structures or by (2) postponing reproduction until the following growing season. We tested these response patterns with the perennial umbellifer Pimpinella saxifraga by simulating flower herbivory and shoot grazing both in the field and in a common garden experiment. In the field, both simulated flower herbivory and grazing effectively suppressed current reproduction, whereas no statistically significant effects of previous-year treatments on growth or reproduction were found in the following year. In the common garden, in the first year the species fully compensated for simulated flower herbivory in vegetative parameters but seed set was reduced by 26%. After 2 years of flower removal, the plants overcompensated in shoot and root biomass by 47 and 46%, respectively, and compensated fully in reproductive performance. Simulated grazing resulted in 21% lower shoot biomass in the first season, but the root biomass was not affected. In the second season the root biomass increased by 43% as compared to the control plants. However, regrowth following simulated grazing resulted in a significant delay in flowering with the consequence that the seed yield of fertile plants was reduced by 55% as compared to the control plants. These results suggest that in resource-rich garden conditions P. saxifraga may immediately repair injuries caused by flower herbivory, but repairs more extensive shoot injury less successfully. Delayed phenology decreases the benefits of immediate repair. In resource-poor conditions, the benefits of regrowth can be negligible. Accordingly, in our field population, the plants postponed their reproduction until the following year in response to simulated grazing and frequently in response to flower removal. When the plants gain very little from regrowth, the costs of reproduction would select for postponed reproduction in response to injury.     

Seed production of cut-leaved teasel (<Emphasis Type="Italic">Dipsacus laciniatus</Emphasis>) in central Missouri

Cut-leaved teasel is an invasive weed in Missouri that reduces the diversification of native species along roadsides and impairs traffic visibility. Teasel is a biennial and grows as a rosette in the first year and flowers the second year. Reproduction is only by seed. Field studies were conducted in 2004 and 2005 at two locations to assess the seed production of cut-leaved teasel. From a natural stand, fifteen plants were tagged at the onset of flowering. Selected plants included those considered growing in a group and those growing alone; a plant was considered alone when no other plant was adjacent for at least 60 cm. Whenever a seedhead completed flowering, it was covered with a cellophane bag and harvested one month later. Linear regression was used to correlate the weight of seeds from a single seedhead and number of seeds to estimate the total seed production per seedhead. The number of seedheads per plant varied from 3 to 56. On average, plants growing alone had 64% more seedheads per plant than plants occurring in a group. Seed numbers in the primary seedhead ranged from 511 to 1,487. Total seed production per plant ranged from 1,309 to 33,527. Seed production was 61% greater for plants growing alone versus those growing in a group and was more prolific in 2005 than in 2004. In addition, seed production per plant varied between locations for plants growing alone, but seed yield per plant was similar for plants growing in groups. Colonization of teasel in new areas is facilitated by higher seedhead numbers per plant and total seed production compared to reproduction of plants in areas of intraspecific competition.     

How do plants know when other plants are flowering? Resource depletion,pollen limitation and mast‐seeding in a perennial wildflower

Mast-seeding is the synchronous and periodic reproduction by plant populations. This phenomenon has been widely studied from a community-level perspective, but we know extremely little about how plants are able to synchronize reproduction. Here, we present the first experimental test of proximate mechanisms of mast-seeding, by preventing reproduction in an iteroparous, mast-seeding wildflower. Through a series of experiments, we show that mobile carbohydrate stores (NSC) control alternate-year flowering by individual plants; seed set depletes NSC which prevents flowering the following year. Plants are synchronized by density-dependent pollen limitation; when plants flower asynchronously, they set fewer seeds, which prevents NSC depletion. Therefore, these individual plants flower in subsequent years and become synchronized. Because mast-seeding is a consequence of physiological controls of reproduction, differences in plant resource acquisition and allocation could dramatically change patterns of seed production, and changes in plant consumers and pollinators could change selection on physiological and developmental pathways.     

Consequences of preformation for dynamic resource allocation by a carnivorous herb,Pinguicula vulgaris (Lentibulariaceae)

When resource availability changes frequently and unpredictably, natural selection favors flexible resource allocation; however, such versatility may be compromised in perennial plants that differentiate leaves or flowers a year in advance of their development (preformation). We investigated resource allocation by the carnivorous perennial Pinguicula vulgaris to determine whether increased resource availability changes within-season allocation to growth, vegetative propagation, and reproduction. In response to resource supplementation (feeding with fruit flies), plants attained a mass 60% greater than that of unfed plants after a single growing season. Feeding also enhanced vegetative propagation, which is closely associated with growth, without modifying relations between these two vegetative functions. In contrast, feeding did not alter the size of vegetative rosettes or the within-season incidence of either flowering or fruiting. This lack of immediate responses occurred because floral and leaf primordia differentiated up to 10 mo before resource supplementation and flower development. However, enhanced resource status likely affected future reproduction indirectly through resource effects on plant size. Large plants produced more floral primordia and between-year changes in fruiting status corresponded to changes in plant size. These results illustrate that preformation can delay responses to enhanced resources by perennial plants.     

Somatic and demographic costs under different temperature regimes in the late-flowering alpine perennial herb Saxifraga stellaris (Saxifragaceae)

If reproduction involves costs, preventing reproduction one year should result in increased growth and/or reproduction the next year. Costs of reproduction in the late-flowering perennial alpine herb Saxifraga stellaris were studied in an experimental field study. To determine whether cost of reproduction differed between two contrasting temperature regimes, I examined plants in Open Top Chambers (OTCs) and control plots. One year before measurements all flower buds on the experiment plants were removed. There was no impact of the flower bud removal or the interaction between flower removal and temperature on prefloration time or seed maturation time. However, cost of reproduction influenced growth (somatic cost), vegetative propagation, flowering frequency, number of stem per ramet, number of fruits, and seed mass (demographic cost). However, significant interaction effects of flower removal and temperature on growth and fruit production revealed that the cost of reproduction differed between the two temperature regimes. The warmed plants showed reduced cost of reproduction compared to plants growing under natural temperature.     

Differences in response to defoliation between males and females of Silene dioica

Summary The response by male and female plants to herbivory was studied by experimental defoliation of the dioecious perennial herb Silene dioica in a green-house. Male and female plants were defoliated prior to and during the early flowering phase at two intensities (50% and 100% of leaf-area removed) in two consecutive years. Defoliation resulted in a decrease in the number of flowers initiated in both sexes, while a larger delay of peak flowering and a higher mortality was observed in males compared to females. In female plants, severe defoliation resulted in a reduction in seed number per capsule and in seed size compared to control. Females showed a negative correlation between the production of flowers in the first and second season in all treatments, while flowering in males the first season was not correlated with flowering in the second season. Females also showed a lower frequency of flowering than males during the two seasons studied. However, during the flowering period, males allocated significantly more biomass to flowers than did females. This outcome supports the idea that females may have a higher total reproductive expenditure than males, but males have a higher reproductive effort during flowering. Male rosette leaves were significantly preferred by the generalist herbivore Arianta arbustorum in experiments. This preference was most pronounced in trials with leaves from fertilized plants compared to nonfertilized plants. A greater storage of resources in aboveground leaves during winter by males compared to females may explain the higher preference for male leaves and the higher male mortality following early defoliation. Furthermore, males are smaller than females and may have a lower ability than females to replace lost resources needed for reproduction when defoliated early in the season.     

Compensation of seed production after severe injury in the short-lived herb Barbarea vulgaris

A pot experiment with the common ruderal herb Barbarea vulgaris (Brassicaceae) was set up to elucidate to what extent short-lived species sprouting from roots regenerate and compensate for seed production after damage. We tested if sprouting from roots ensures survival after severe aboveground biomass damage, but the number of seeds produced declines with increasing severity of injury, decreasing nutrient availability and progress in the life cycle at the time of injury.Plants of B. vulgaris were cultivated in a 3-year garden experiment at two nutrient levels (high vs. low). During the experiment, two levels of injury severity were applied: high (removal of all aboveground biomass) and low (removal of aboveground biomass leaving basal axillary buds intact). Damage was applied at four life-cycle phases: young rosette, overwintered rosette, flowering plant and fruiting plant. All injured plants survived and resprouted irrespective of life-cycle phase, severity of injury and nutrient availability. Injury significantly affected seed production and also the plants’ life cycle. Plants injured in the second year of their life (overwintered rosette, flowering plant and fruiting plant) postponed reproduction to the third season (in the case of high injury severity) or their seed production was lower than in intact plants (in the case of low injury severity). In plants injured in the first life year, seed production and life cycle were not influenced. Nutrient level only marginally affected resprouting after injury and seed production.The experiment showed that the ability to sprout from roots enables plants to survive a 100% loss of aboveground biomass, and to keep some seed production or even compensate it. The short-lived ruderal species B. vulgaris successfully copes with severe disturbance by resprouting and does not rely only on its seed bank.     

Effects of fire season on flowering of forbs and shrubs in longleaf pine forests

Summary Effects of variation in fire season on flowering of forbs and shrubs were studied experimentally in two longleaf pine forest habitats in northern Florida, USA. Large, replicated plots were burned at different times of the year, and flowering on each plot was measured over the twelve months following fire. While fire season had little effect on the number of species flowering during the year following fire, fires during the growing season decreased average flowering duration per species and increased synchronization of peak flowering times within species relative to fires between growing seasons. Fires during the growing season also increased the dominance of fall flowering forbs and delayed peak fall flowering. Differences in flowering resulting from variation in fire season were related to seasonal changes in the morphology of clonal forbs, especially fall-flowering composites. Community level differences in flowering phenologies indicated that timing of fire relative to environmental cues that induced flowering was important in determining flowering synchrony among species within the ground cover of longleaf pine forests. Differences in fire season produced qualitatively similar effects on flowering phenologies in both habitats, indicating plant responses to variation in the timing of fires were not habitat specific.     

Selection on flowering time in Mediterranean high-mountain plants under global warming

Under climate warming, plants will undergo novel selective pressures to adjust reproductive timing. Adjustment between reproductive phenology and environment is expected to be higher in arctic and alpine habitats because the growing season is considerably short. As early- and late-flowering species reproduce under very different environmental conditions, selective pressures on flowering phenology and potential effects of climate change are likely to differ between them. However, there is no agreement on the magnitude of the benefits and costs of early- vs. late-flowering species under a global warming scenario. In spite of its relevance, phenotypic selection on flowering phenology has rarely been explored in alpine plants and never in Mediterranean high mountain species, where selective pressures are very different due to the summer drought imposed over the short growth season. We hypothesized that late-flowering plants in Mediterranean mountains should present stronger selective pressures towards early onset of reproduction than early-flowering species, because less water is available in the soil as growing season progresses. We performed selection analyses on flowering onset and duration in two high mountain species of contrasting phenology. Since phenotypic selection can be highly context-dependent, we studied several populations of each species for 2 years, covering their local altitudinal ranges and their different microhabitats. Surrogates of biotic selective agents, like fruitset for pollinators and flower and fruit loss for flower and seed predators, were included in the analysis. Differences between the early- and the late-flowering species were less than expected. A consistent negative correlational selection of flowering onset and duration was found affecting plant fitness, i.e., plants that bloomed earlier flowered for longer periods improving plant fitness. Nevertheless, the late-flowering species may experience higher risks under climate warming because in extremely warm and dry years the earlier season does not bring about a longer flowering duration due to summer drought.     

Sex ratio variation in dioecious plant species: A comparative ecological study of six species of Lindera (Lauraceae)

Sex ratio variation was investigated in natural populations of six dioecious shrub species of Lindera in Japan. Interspecific differences in sex ratio were examined in relation to patterns of population structure, floral dimorphism, fruit production and intersexual differences in herbivory. Sex ratios tended towards equality or bias in favour of males, except for populations of L. glauca , in which no male plants were found. Sex switching in individual plants was not observed. Although male flowers were generally larger in size than flowers of conspecific females, costs of flowering did not appear to influence sex ratio. However, a relation was found between high allocation to fruit production and increasing male fraction of populations. In addition, girths of flowering plants tended to be larger in females, which may indicate delay in age of the first reproduction compared to males. No intersexual difference was found in degree of leaf damage due to herbivory. Results of this study are discussed in the light of other studies on sex ratio variation in dioecious plant species.     

Optimal phenology of annual plants under grazing pressure

Plants show phenological responses to herbivory. Some enclosure experiments have demonstrated that the onset of the peak flowering season is dependent on grazing pressure. We constructed a mathematical model using Pontryargin's maximum principle to investigate changes in flowering time by examining shifts in resource allocation from vegetative to reproductive plant components. We represented a primary production of a plant individual by two types of function of vegetative part size, a linear function and a convex non-linear function. The results of a linear production model indicate that optimal phenology follows a schedule that switches from the production of vegetative parts to that of reproductive parts at a given time ('bang-bang' control). However, in a non-linear model, a singular control, wherein the plant invests in both productive and reproductive parts, may be included between obligate production and reproduction periods. We assumed that the peak of the flowering season occurs immediately following the exclusive investment in reproduction. In a linear production model, differential herbivory rates on the vegetative and reproductive parts of a plant resulted in shifts in the peak flowering time. A higher herbivory rate on the vegetative components advanced the peak, whereas it was delayed when grazing pressure focused on reproductive components of the plant. In the non-linear production model, increased grazing pressure tended to postpone the flowering peak. These results corresponded well with results of enclosure experiments, thus suggesting adaptive control of flowering time in plants.     

Production dependence of vegetative propagation inDisporum smilacinum A. Gray

The dependence of vegetative propagation on the production of individual plants was examined inDisporum smilacinum A. Gray on the basis of shading experiments and field surveys. This species typically showed four types of reproductive behavior: sterile plants producing one plantlet (no propagation), sterile plants producing more than two plantlets (vegetative propagation), fertile plants producing one plantlet (sexual reproduction) and fertile plants producing more than two plantlets (both sexual reproduction and vegetative propagation). The propagation ofD. smilacinum was clearly related to the annual net production of each individual plant. The probability of a mother plant producing more than two vegetative propagules (plantlets) increased with net production of the plant in the current year. The number of propagules per plant and runner length increased with net production. It was possible to explain the types of reproductive behavior of this species on the basis of both the initial plant size before sprouting and its net production during the growing season. There was a critical initial plant size for sexual reproduction and a critical level of production for vegetative propagation.     

Flowering phenology and reproduction of a forest understorey plant species in response to the local environment

Temperate forest understorey plants are subjected to a strong seasonality in their optimal growing conditions. In winter and early spring, low temperatures are suboptimal for plant growth while light becomes limited later in spring season. We can thus expect that differences in plant phenology in relation to spatiotemporal environmental variation will lead to differences in reproductive output, and hence selection. We specifically studied whether early flowering, a paradoxical pattern that is observed in many plant species, is an adaptive strategy, and whether selection for early flowering was confounded with selection for flower duration or was attributable to environmental variables. We used Geum urbanum as a study species to investigate the effect of relevant environmental factors on the species’ flowering phenology and the consequences for plant reproductive output. We monitored the phenology of four to six plants in each of ten locations in a temperate deciduous forest (Belgium). We first quantified variation in flowering time within individuals and related this temporal variation to individual flower reproductive output. Then, we studied inter-individual variation here-in and linked this to reproduction at the plant level, hence studying the selection differential. We found that flowering within individual plants of Geum urbanum was spread over a long period from June to October. Reproductive output of individual flowers, measured as total seed mass per flower, declined during the season. We found no indication for selection for early flowering but rather for longer flower duration. Larger plants had an earlier flowering onset and a higher seed mass, which suggests that these factors covary and are condition dependent. None of the studied environmental variables could explain plant size, although soil pH and to a lesser extent light availability had a positive direct effect on seed mass per plant. Finally, we suggest that the high intra-individual variation in flowering time, which might be a risk spreading strategy of the plant in the presence of seed predation, limits the potential for selection on flowering phenology.

     

Differential costs of reproduction in females and hermaphrodites in a gynodioecious plant

Background and Aims

Plants exhibit a variety of reproductive systems where unisexual (females or males) morphs coexist with hermaphrodites. The maintenance of dimorphic and polymorphic reproductive systems may be problematic. For example, to coexist with hermaphrodites the females of gynodioecious species have to compensate for the lack of male function. In our study species, Geranium sylvaticum, a perennial gynodioecious herb, the relative seed fitness advantage of females varies significantly between years within populations as well as among populations. Differences in reproductive investment between females and hermaphrodites may lead to differences in future survival, growth and reproductive success, i.e. to differential costs of reproduction. Since females of this species produce more seeds, higher costs of reproduction in females than in hermaphrodites were expected. Due to the higher costs of reproduction, the yearly variation in reproductive output of females might be more pronounced than that of hermaphrodites.

Methods

Using supplemental hand-pollination of females and hermaphrodites of G. sylvaticum we examined if increased reproductive output leads to differential costs of reproduction in terms of survival, probability of flowering, and seed production in the following year.

Key Results

Experimentally increased reproductive output had differential effects on the reproduction of females and hermaphrodites. In hermaphrodites, the probability of flowering decreased significantly in the following year, whereas in females the costs were expressed in terms of decreased future seed production.

Conclusions

When combining the probability of flowering and seed production per plant to estimate the multiplicative change in fitness, female plants showed a 56 % and hermaphrodites showed a 39 % decrease in fitness due to experimentally increased reproduction. Therefore, in total, female plants seem to be more sensitive to the cost of reproduction in terms of seed fitness than hermaphrodites.     

Effect of flowering on vegetative growth and further reproduction in

Flowering intensity and plant size were monitored in 155 Festuca novae-zelandiae individuals over four years to determine if trade-offs exist between inflorescence production and vegetative growth, and between inflorescence production in different years. Less than half of the population flowered in any one year, 36% of individuals did not flower at all, and only 17% flowered in all four years of the study. Mean number of inflorescences per individual per year varied from 1.54 to 5.53 (maximum = 85). No trade-offs were detected between flowering frequency and intensity; individuals that flowered more frequently also produced more inflorescences in each flowering episode. No trade-off was detected between current and future reproduction, rather flowering intensity was positively correlated between years. Growth, as measured by diameter increment, was positively related to flowering frequency and flowering intensity, both across all individuals studied and within 1m x 1m plots. The presence of a positive relationship between growth and reproduction within plots argues against meso-scale variability in environment factors being the cause of the results from analyses involving all individuals. Clearly reproduction in F. novae-zelandiae does not incur a marked cost in growth or future reproduction. The assumptions underlying theoretical expectations of such trade- offs may not be valid for long-lived clonal plants such as F. novae-zelandiae.     

Costs of reproduction in Nyssa sylvatica: sexual dimorphism in reproductive frequency and nutrient flux

Summary We examined the influence of differential reproductive frequency between the sexes on tertiary (phenotypic) sex ratios in the the dioecious tree Nyssa sylvatica (Nyssaceae). Reproduction was evaluated in relation to sex, size and canopy exposure using flowering data collected from 1229 marked trees over a four year period. For subsets of each population we used data on flower number, fruit crop size, fruit/flower ratios, and individual flower and fruit mass to compare biomass invested in reproductive structures of males and females. We also examined seasonal changes in stem nitrogen and soluble carbohydrate content in relation to flower and fruit production for trees of each sex. Our results indicate that: 1) Male-biased tertiary sex ratios could be explained by more frequent reproduction by male trees; 2) Estimated secondary sex ratios based on sums of all known males and females were not significantly different from 1:1; 3) Flowering frequency of males and females was significantly related to plant size (DBH) and exposure of the canopy to light; 4) Estimtes of reproductive biomass allocation ranged from 1.36 to 10.8 times greater for females relative to males; 5) Flower production was related to stem nutrient status for both sexes, but nutrient depletion and its effect on subsequent flowering was much more pronounced for female trees. We conclude that less frequent flowering by female trees may result from depletion of stored reserves, and that differential flowering frequency in N. sylvatica may ultimately reduce apparent sexual differences in the costs of reproduction.