Reproduction in an Orchid Can Be Resource‐Limited over its Lifetime1

Most orchids studied thus far show long‐term resource adjustments to increases in fruit production within a flowering season, but none of these offers rewards to their potential pollinators. If nectar production is energetically expensive, then resources utilized to produce fruits and seeds may be even more limited in pollinator‐rewarding orchids than in non‐rewarding ones. Thus, resource adjustments may be more dramatic or entirely different in nectar producing plants. In this study, we performed artificial hand‐pollinations for two consecutive flowering seasons in die nectar producing orchid Comparettia falcata, and tested whether or not fruit set, seed set, and seed viability were limited by the quantity of pollinations or by resources. In addition, we compared mechanisms of short‐term (fruit abortion within seasons) and long‐term consequences (percent change in leaf length and change in flower number per plant between seasons, probability of shoot and inflorescence production, and mortality) between hand‐ pollinated and unmanipulated plants. The relationships among plant traits related to vegetative size and reproduction also were examined. Hand‐pollinations showed some negative effects. Fruit set was higher in hand‐pollinated plants in the first season but was similar to the controls in the second. Seed set was significantly lower and abortions were higher than in unmanipulated plants. On the other hand, some of our measurements were unaffected by die hand‐pollination treatment. Unexpectedly, there were no significant differences between groups in percent change in leaf length, change in flower number per plant between seasons, or die probability of shoot and inflorescence production. Although there was a strong correlation between leaf size and die number of flowers produced within a season, associations between leaf size and traits related to current or future reproduction were not consistent. Like other epiphytic orchids, pollination limitation occurred within a single season in C. falcata., but increases in fruit production also resulted in reduced lifetime fitness as estimated by a compounded fitness index. Contrary to all other epiphytic orchids studied, long‐term adjustments to increased fruit production in C. falcata through reduction in future growth or flower and inflorescence production were either minor or lacking. Our results suggest that the nature of plant strategies associated with resource constraints during sexual reproduction may be dependent on whether or not plants have evolved traits that are costly.     

FOUR-YEAR GROWTH AND REPRODUCTION OF CYCLOPOGON CRANICHOIDES (ORCHIDACEAE) IN SOUTH FLORIDA

Thirty-five flowering individuals of the orchid Cyclopogon cranichoides were marked in 1986, and their leaf area, number of flowers, and number of fruits were recorded from 1986 to 1989. In 1986 and 1987 all the flowers on 20 individuals were hand-pollinated. The number of plants with above-ground growth steadily decreased by about 20% each year. In some cases there was evidence that disappearance of plants was not due to mortality but to a transition into a subterranean state. Small plants were most likely to make this transition. Flower number was significantly associated with plant size. Fruiting success was pollinator-limited throughout this study; fruit set of hand-pollinated individuals was higher than 96%, whereas natural fruit set ranged from 26% in 1987 to 62% in 1988. Fruit set did not have any effect on growth or reproduction the following year.     

The Effect of Rust Infection on Reproduction in a Tropical Tree (Faramea occidentalis)1

Fungal pathogens that infect reproductive structures of plants (e.g., flowers and fruits) can reduce the seed production and seedling recruitment of host plants. We report here on the effects of a rust, Aecidium farameae, that infects the ovaries and pedicels of mature flowers on Faramea occidentalis (Ruhiaceae), a small tree common on Barro Colorado Island, Republic of Panama. Rust infection of ovaries reduced the number of maturing fruit on infected trees. Trees with low rust incidence in June of 1992 had 68 percent fruit survival, compared to 17 percent fruit survival for those with high rust incidence. Infected fruits developed abnormally and were usually aborted long before uninfected fruits were mature. One hundred percent of the infected ovaries marked in July were diseased or missing in August. We conclude that infection by A. farameae has the potential to seriously decrease the reproductive output of Faramea occidentalis and may represent an important source of variation in the relative fitness of individual plants.     

Density-dependent variation in reproductive success in a terrestrial orchid

 Within population variation in plant density can affect reproductive success and breeding systems. We examined such effects in Listera cordata R. Br., a north temperate terrestrial orchid, in a subalpine population in Colorado, USA. Hand pollinations showed that L. cordata was self-compatible as all pollinations produced fruits. Cross-pollinations differed significantly from self-pollinations, and had a higher number of seeds per capsule and higher % of fertilized ovules. Together they could compensate for the transmission advantage of self-pollinations. Average number of flowers per inflorescence was 4.2–4.4 and average fruit set was 20.7–23.7%. By establishing 36 plots with varying plant densities (1, 4, 8 and 16) and emasculating all flowers in target plants of a set of 20 plots, we estimated density effects on fruit and seed production. Plant density had no effect on the amount of reproduction except in reduced seed production of highly dispersed plants. Our results suggest that inbreeding depression may be an important selective factor in L. cordata. Further studies are needed to define its role in the evolution of mating systems in this species. Received February 21, 2000 Accepted December 26, 2000     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Effects of mycorrhizal infection, soil phosphorus availability and fruit production on the male function in two cultivars of Lycopersicon esculentum

The effects of mycorrhizal infection, soil P availability and fruit production on the male function of reproduction were examined in two cultivars of tomato (Lycopersicon esculentum Mill.). Tomato plants were grown in a greenhouse under three treatment combinations: non‐mycorrhizal, low P (NMPO); non‐mycorrhizal, high P (NMP3); and mycorrhizal, low P (MPO). In addition, all treatment combinations were grown both with and without fruit. Fruit production decreased final leaf biomass, flower production and in vitro pollen tube growth rates, often reducing the beneficial effects of increased P uptake. Thus, fruit production diverted resources from subsequent vegetative growth, flower production and pollen development. As the growing season progressed, mean pollen production per flower and in vitro germination and tube growth decreased. Mycorrhizal infection and high soil P conditions increased final leaf biomass, flower production, mean pollen production per flower (in one cultivar) and in vitro pollen tube growth rates. Thus, mycorrhizal infection and high soil P conditions increased pollen quantity and quality, thereby enhancing fitness through the male function. Similar trends in these treatments suggested that mycorrhizal effects on the male function were largely the result of improved P acquisition.     

Costs of reproduction in the pink lady's slipper orchid (Cypripedium acaule): defoliation,increased fruit production,and fire

An earlier experiment with the pink lady's slipper orchid demonstrated that plant leaf area was lowered only after successive years of increased fruit production. This result suggested that the cost of reproduction was small in relation to the energy budget of the plant. To test this idea, plants were subjected to experimental hand-pollination treatments to increase fruit set as well as leaf removal treatments to decrease the energy budget of plants. Changes in plant size in years 2 and 3 and, to some extent, rate of flowering, were determined by a combination of initial plant size, leaf removal treatments in year 1, fruit production in year 1, and damage from an unplanned fire in year 2. Plants that had both leaves removed and produced a fruit in 1987 decreased in size in the following 2 years in comparison with other treatment groups. The cost of fruit production was not apparent in plants that had only one or no leaves removed. Plants apparently have to be put into severe physiological stress in order for a cost of reproduction to appear in the following year. The cost of producing one fruit was a decline of plant size in the following year of 30 cm², which is very similar to our previous experiment using a different design. An additional experiment failed to find evidence that these plants increase their photosynthetic rate to compensate for the loss of leaves or the cost of maturing fruit. Published experiments in both the greenhouse and the field that failed to find a cost of reproduction should be reevaluated in terms of the intensity of treatment imposed and the overall energy budget of the plant in field situations.     

Regulation of reproduction in wild and cultivatedLycopersicon esculentum Mill. by vesicular-arbuscular mycorrhizal infection

Summary We examined how mycorrhizal infection byGlomus etunicatum Becker and Gerd. affected flowering phenology and components of reproduction in eight wild accessions and two cultivars ofLycopersicon esculentum Mill. We did this by performing a detailed demographic study of flower, fruit and seed production. Mycorrhizal infection had variable effects on the ten accessions. Infection significantly decreased the time taken to initiate flowering in some accessions. In addition, infection increased flowering duration in some accessions. In many accessions, infection significantly increased seed production, primarily by increasing the number of inflorescences and infructescences. In some accessions, mycorrhizal infection also increased the proportion of flowers that produced mature fruits or the number of seeds per fruit. Among accessions, shoot phosphorus content was correlated with seed productivity for both mycorrhizal and non-mycorrhizal plants. However, non-mycorrhizal plants produced more seed biomass per mg of shoot phosphorus than mycorrhizal plants.     

Effects of Aphis fabae and Uromyces viciae-favae on the growth of a susceptible and an aphid resistant cultivar of Vicia faba

Damaging effects of either black bean aphid (Aphis fabae), broad bean rust (Uromyces viciae-fabae), or the combination of both were investigated on a susceptible (cv. Diana) and an aphid resistant (cv. Bolero) cultivar of Vicia faba. When compared with rust, aphids caused greater reductions of root dry weight, shoot dry weight, leaf area, and mean relative growth rate. The mean unit leaf rate was also reduced whereas the leaf area ratio was not affected. The damage caused per aphid was highest on the susceptible cultivar. Rust induced damage did not differ between the cultivars. Concomitant infestation with both pests only resulted in additive damage. The population development of aphids was delayed on partially resistant plants. High temperature and rust infection reduced the total number of aphids the plants were able to support but not the level of resistance. Thus the specific damaging effect per aphid was increased.     

Host-pathogen interactions in natural populations of Linum marginale and Melampsora lini

Summary The epidemiology of rust caused by the fungus Melampsora lini and the effects of infection by this pathogen on its host, the herbaceous perennial Linum marginale, were determined in the field and in garden experiments. There was considerable natural variability in disease levels over the four years (1986–1989) of the study. In two years (1986, 1989) major rust epidemics occurred. In the field, the main effect of disease was to reduce survivorship during the winter following infection. Plants which were heavily infected during the 1986 or 1989 growing seasons had reduced survivorship relative to more lightly infected plants. Melampsora lini infections did not appear to affect survivorship in either 1987 or 1988. Flowering was correlated with environmental factors and the number of stems a plant possessed. A severe drought in 1987 completely inhibited flowering. In the other three years the number of flowers produced by a plant was strongly positively correlated with the number of stems it possessed. Disease levels had no consistent effect on flowering. Controlled garden experiments were also used to examine the response of seedlings and adult plants to infection. These showed that both the timing and severity of disease appears to determine the effect of M. lini infections on L. marginale. Early, severe infection reduced growing season and overwintering survivorship as well as capsule production. However, plants in the field were most often infected only after flowering had begun, and the predominant effect of infection was a reduction in overwintering survivorship. The high variability in disease levels from year to year and the deferred nature of the effect of the rust on its host have significant implications for the design of experiments aimed at assessing the role of diseases in plant communities.     

Fruit set in a deceptive orchid: The effect of flowering phenology,display size,and local floral abundance

We investigated the effect of flowering time, display size, and local floral density on fruit set in Tolumnia variegata, a pollination-limited orchid that offers no reward to its pollinator(s). During 1990, natural variation in flowering time, display size, and fruit set were monitored in 508 plants at one locality in Puerto Rico. The following season, orchid floral abundance per host tree (Randia aculeata) was manipulated to investigate its effect on fruit set. Four floral abundance treatments were established (700, 500, 300, and 100), each replicated four times. Flowering time was the most important trait affecting fruit set. The proportion of plants setting at least one fruit was significantly high early and late in the season, but low during the flowering peak. Thus, strong disruptive selection differential on flowering phenology was found. Display size had little effect on fruit set. A weak, but significant disruptive selection differential on display size was found. Orchid floral abundance per host tree had a significant effect on fruit set. Early in the season, T. variegata flowers with intermediate number of conspecific flowers exhibited a greater probability of setting fruit than those in host trees with fewer or more flowers. Our results show that flowering phenology may be evolutionarily unstable, possibly a consequence of the deception pollination system. Furthermore, a deception strategy would be relatively unsuccessful in populations where plants are found in either very dense or sparse patches.     

Cost of reproduction in the pink lady's slipper orchid (Cypripedium acaule, Orchidaceae): an eleven-year experimental study of three populations

An 11-yr experimental study of the cost of reproduction in three wild populations of the perennial orchid Cypripedium acaule contrasted experimental plants that were repeatedly hand-pollinated and often made fruits with control plants that were not hand-pollinated and only rarely made fruits. Repeated flowering without subsequent fruit production resulted in no detectable reduction in either plant size or probability of flowering in subsequent years. A cost of fruit production was evident in experimental plants in all three populations in terms of a reduced probability of flowering and smaller leaf area in subsequent years, but was not evident in terms of mortality rate. Experimental effects of fruit production reached maximum values at 3-7 yr, depending on the population. The probability of remaining dormant below ground in a given year was strongly dependent on plant size in the previous year. Furthermore, the length of the dormancy period (one to several years) was a significant and inverse function of plant size just prior to dormancy. Sample sizes and the consequent ability to detect experimental effects declined over time as more plants died or stopped flowering. Four to seven years appears to be an optimal duration for studies of the cost of reproduction in perennial herbs similar to this species. Studies lasting less than 4 yr may be too brief to reveal experimental effects, whereas those lasting more than 7 yr may fail to reveal new insights.     

Nitrogen-induced reduction in leaf phenolic level is not accompanied by increased rust frequency in a compatible willow (Salix myrsinifolia)‐Melampsora rust interaction

Phenolic compounds are potential antimicrobial agents in willows ( Salix spp.). However, little is known about their role in willow‐pathogen interactions. Earlier studies have shown that phenolic levels may vary depending on genotype and environmental factors (such as nutrient availability). In the present study, the combined effects of nitrogen fertilization and infection by pathogenic rust ( Melampsora sp.) on phenolics and growth in willow ( salix myrsinifolia ) clones were investigated. Cutting-propagated plants of eight willow clones were subjected to low and optimum levels of nitrogen fertilization and inoculated with rust urediniospores or mock-inoculated with water. The plants were harvested 20 days after the inoculations and the dry weight of the leaves, stems and roots determined. Phenolics were analysed from leaf material using HPLC. The willow clones differed significantly in their phenolic content, rust frequency and phytomass production. Plants under optimum nitrogen availability showed increased shoot phytomass production and reduced root production. Rust infection increased the root phytomass of willow under both fertilization treatments, but had no consistent effect on the shoot growth of the clones. Low nitrogen availability increased the phenolic levels in willow leaves, but the rust frequency did not differ significantly between plants grown at low and optimum levels of nitrogen fertilization. In several clones, rust infection increased the concentrations of individual phenolic compounds, but there were no significant correlations between phenolic concentrations and the level of rust infection. The results indicate that resistance in compatible willow‐rust interactions may not be directly governed by phenolic compounds.     

Effects of above- and belowground herbivory on growth,pollination, and reproduction in cucumber

Plants experience unique challenges due to simultaneous life in two spheres, above- and belowground. Interactions with other organisms on one side of the soil surface may have impacts that extend across this boundary. Although our understanding of plant–herbivore interactions is derived largely from studies of leaf herbivory, belowground root herbivores may affect plant fitness directly or by altering interactions with other organisms, such as pollinators. In this study, we investigated the effects of leaf herbivory, root herbivory, and pollination on plant growth, subsequent leaf herbivory, flower production, pollinator attraction, and reproduction in cucumber (Cucumis sativus). We manipulated leaf and root herbivory with striped cucumber beetle (Acalymma vittatum) adults and larvae, respectively, and manipulated pollination with supplemental pollen. Both enhanced leaf and root herbivory reduced plant growth, and leaf herbivory reduced subsequent leaf damage. Plants with enhanced root herbivory produced 35% fewer female flowers, while leaf herbivory had no effect on flower production. While leaf herbivory reduced the time that honey bees spent probing flowers by 29%, probing times on root-damaged plants were over twice as long as those on control plants. Root herbivory increased pollen limitation for seed production in spite of increased honey bee preference for plants with root damage. Leaf damage and hand-pollination treatments had no effect on fruit production, but plants with enhanced root damage produced 38% fewer fruits that were 25% lighter than those on control plants. Despite the positive effect of belowground damage on honey bee visitation, root herbivory had a stronger negative effect on plant reproduction than leaf herbivory. These results demonstrate that the often-overlooked effects of belowground herbivores may have profound effects on plant performance.     

兰科菌根真菌研究方法的概述

兰科植物资源在全球分布广泛,其中有许多是重要的药用植物和名贵的珍稀花卉,由于具有较高的商业价值,受到各界人士的广泛关注。兰科植物生长习性的特殊性导致其在自然状态下繁殖率极低,因此难以满足市场的广泛需求。近年研究表明,几乎所有兰科植物都能与相应的菌根真菌建立共生关系,并且必须依赖于这些内生真菌才能完成其整个生活史。因而对菌根真菌在提高兰科植物生长速度和繁殖能力过程中机制的研究以及将研究成果运用于工业化育苗中将是缓解兰科植物市场供求紧张问题的关键。通过对近几年有关天麻和铁皮石斛等兰科植物的问题研究中所采用的研究方法加以阐述,以期对今后兰科菌根真菌的研究提供一定的参考。     

Sexual dimorphism and cost of reproduction in the dioecious shrub Lindera benzoin (Lauraceae)>

We examined sexual dimorphism in reproductive allocation and its effects on growth and subsequent reproduction in a natural population of the dioecious woody shrub Lindera benzoin (L.) Blume. In addition to comparing natural patterns of growth and reproduction in a marked population of 251 females and 87 males, we experimentally examined the effect of reduced reproduction on future growth and reproduction in female plants and examined sexual dimorphism in carbon/nutrient balance. Our results suggest that females of L. benzoin bear greater reproductive costs in terms of both biomass and minerals. These costs were measurable in terms of current biomass and allocation of nitrogen to reproduction, as well as subsequent growth, reproduction, and tissue carbon/nutrient balance. Based upon the results of a fruit-thinning experiment and path analysis, fruit production in 1991 appeared to have direct negative effects on fruit production in 1992, an effect that was not necessarily mediated through effects on plant growth. We discuss our results in the context of other intrinsic and extrinsic factors that can influence growth, reproductive costs, and mortality in this species.     

Pollen limitation and fruiting failure related to canopy closure in Calypso bulbosa (Orchidaceae), a northern food‐deceptive orchid with a single flower

Natural fruit set is constrained by pollen limitation and fruiting failure, and pollen limitation is expected to be especially severe in deceptive orchids. We performed hand cross‐pollinations in ten populations of a food‐deceptive orchid, Calypso bulbosa, under sparse and dense canopies in three non‐consecutive years. We explored the relationships between natural fruit set, pollen limitation and fruiting failure. Mean natural fruit set over the years was 60%, which is exceptionally high for a deceptive orchid. On average, hand cross‐pollination increased fruit set by 23%. Among open‐pollinated plants that did not set a fruit, 55.5% were estimated to be pollen limited and 44.5% to be limited by fruiting failure, i.e. inability to set a fruit after pollination. In species with high natural fruit set, hand cross‐pollination experiments may not always detect statistically significant pollen limitation. In our case, pollen limitation tended to become significant when the natural fruit set dropped below 60%. Canopy cover had a significant effect on fruiting failure, which was more severe under a dense canopy. Although our results demonstrate pollen limitation in many cases, they also highlight the fact that food deception can be a very effective pollination strategy. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 744–750.     

The Effects of Brown Rust and Phosphate on Cold Acclimation in Winter Barley

The green leaf area of winter barley, cv. Sonja, sampled fromthe field at different times during winter was always greatestin plants grown at high soil phosphate and smallest in plantsgrown at low soil phosphate, and at each fertilizer level wasgreater in healthy plants than in plants infected by rust (Pucciniahordei). In leaves that survived the coldest period of winter,the percentage area that was damaged was increased by rust infectionwhich prevented the ameliorating effects of high soil P. Rustand low P interacted to reduce the increases in leaf area andshoot d. wt that occurred when higher temperatures prevailedin spring. Under controlled conditions in the laboratory, phosphate reducedthe injury suffered when plants not acclimated to low temperatureswere exposed to freezing conditions, but this effect was removedby rust infection. After rust infection, freezing temperatureswere damaging even to acclimated plants, particularly if grownwith low soil P. Evidence of visible symptoms, and quantitativemeasurements of electrolyte efflux from intact leaves, chlorophyllfluorescence in vivo, and ethane and ethylene evolution fromcold-acclimated plants, showed that infection raised the minimumtemperature at which tissues could survive without injury. Infectedleaves were more sensitive to low temperature post-sporulationthan presporulation. Measurements of electrolyte efflux andchlorophyll fluorescence on plants growing under cold conditionsshowed that infection inhibited the processes of acclimationto low temperatures. Winter barley, Puccinia hordei, injury, low temperature, acclimation     

Intercellular chitinase and peroxidase activities associated with resistance conferred by geneLr35to leaf rust of wheat

The effect of leaf rust (Puccinia triticina) infection on intercellular chitinase (EC 3.2.1.14) and peroxidase (EC 1.11.1.7) activities was studied in resistant [RL 6082 (Thatcher/Lr35)] and susceptible (Thatcher) near isogenic wheat (Triticum aestivum L.) lines at seedling, stem elongation and flag leaf stages of plant growth. The levels of activity of these enzymes were low during the seedling and stem elongation stages. Resistant plants at the flag leaf stage, during which the Lr35 resistance gene was maximally expressed, exhibited high constitutive levels of chitinase and peroxidase activities, in contrast to the lower constitutive levels of susceptible plants. The results suggest that chitinase and peroxidase, constitutively present in the intercellular spaces of Thatcher/Lr35 wheat leaves, may play a role in Lr35 mediated resistance to leaf rust.