Population ecology in a natural<Emphasis Type="Italic">syneilesis palmata</Emphasis> stand: II. vegetative growth and population structure

Vegetative growth and changes within populations ofSyneilesis palmata (THUNB>) MAX. Usannamul were monitored in two natural stands with different levels of incoming light. This perennial, shade-tolerant herb was measured for its plant size, petiole length, leaf breadth, and leaflet number over 11 years in a moderately shaded (MS) stand and over 7 years in a severely shaded (SS) stand. At the end of each growing season, a shoot primordium developed at the center of the root system. Correlation coefficients were mostly high between pairings of two properties - total weight, petiole length, petiole weight, leaf breadth, leaf blade weight, leaf area, specific leaf area, and leaflet number. Mean annual mortality of this species was higher in SS (22.97%) than in MS (8.85%), but great fluctuations were seen from year by year. Mortality was lowest for medium-sized plants. Regarding petiole length, differences in mean annual growth rates were conspicuous, i.e., 2.6% in MS and 8.4% in SS, while growth rates for leaf breadth were 3.1% in MS and 24.2% in SS. Changes in plant frequency within individual size classes roughly showed a normal distribution curve. However, the mode varied year by year, and such changes were more remarkable in SS than in MS. Under severe shading,S. palmata had a higher growth rate and but also greater mortality than did plants under moderate shade. Therefore, one can conclude that the population structure in SS was unstable over time.     

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.     

Flowering Frequency and Plant Performance and their Relation to Age in the Perennial Orchid Spiranthes spiralis (L.) Chevall.

Abstract: Long-term demographic data have been analyzed to establish possible costs of flowering in the terrestrial orchid Spiranthes spiralis (L.) Chevall. in The Netherlands. Costs of flowering can be expressed as individual plant performance and flowering frequency in relation to the generative or vegetative status in the following year. Flowering in individuals of S. spiralis in a given year (t) is followed by a non-flowering phase in the next growing season (t + 1) in more than 80 % of the plants. The decline in flowering frequency is not a result of the age structure of the population involved because individual plants do not show signs of senescence after 10 - 15 years of aerial presence as an autotrophic plant. Rosettes have a smaller leaf area in the year of flowering (t), compared to the previous (t - 1) and following year (t + 1), due to the allocation of the limited underground resources to both flowering stalk and rosette at the beginning of its growing season. Generative reproduction in S. spiralis has a significant negative impact on both flowering frequency in subsequent years and on rosette size in the year of flowering. The flowering frequency and rosette size in relation to the life history, characterized by the yearly replacement of the underground tuber, is discussed. Better understanding of the life-history strategy, including costs of reproduction, may contribute to the creation of sustainable environmental conditions for growth of S. spiralis, e.g., optimal conditions for photosynthesis during the aboveground stage of the tiny wintergreen rosettes.     

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.     

Population dynamics ofHeloniopsis orientalis C. Tanaka (Liliaceae) in natural forests — Sexual reproduction

This study investigated the effects of climate on reproduction, consecutive flowering, and the level of energy investment toward sexual reproduction, as well as the relationships between flowering rates and plant size, and between seed production and plant size. The target species wasHeloniopsis orientalis C. Tanaka (Liliaceae), an evergreen perennial herbaceous plant. Natural populations were monitored in permanent quadrats for seven years in a low-elevation area (Namhansanseong) and for five years in an area with a high elevation (Maranggol). The Tn indices required for flowering were relatively similar each year, but those for seed dispersal fluctuated. The period for seed production was proportional to the total length of the growing season. Rates of flowering varied annually, and depended on conditions in each study area. Flowering was discontinuous (i.e., not consecutive) at Namhansanseong, but was continuous (i.e., consecutive) at Maranggol. The rates were proportional to plant size; the minimum size class that supported flowering was 60–90 cm², and almost all plants with >210 cm² leaf area flowered. The overlapping range in plant size between flowering and non-flowering plants was relatively broad. Numbers of fruit per plant ranged from 1-12, with an average of 6.0 fruits at Namhansanseong and 5.0 fruits at Maranggol. In the latter area, the range in the number of seeds produced and their weight per plant varied drastically, being 1700–1867 seeds and 133.2–177.5 mg, respectively. Almost all correlation coefficients between plant size and fruit number, plant size and seed number, or plant size and seed weight were significant at the 1% level. Lastly, energy investment rates for sexual reproduction were about 30–50% of the net productivity, varying over the years. These rates were proportional to growth rates.     

Attracting pollinators and avoiding herbivores: insects influence plant traits within and across years

Perennial plants interact with herbivores and pollinators across multiple growing seasons, and thus may respond to herbivores and pollinators both within and across years. Joint effects of herbivores and pollinators influence plant traits, but while some of the potential interactions among herbivory, pollination, plant size, and plant reproductive traits have been well studied, others are poorly understood. This is particularly true for perennial plants where effects of herbivores and pollinators may manifest across years. Here, we describe two experiments addressing the reciprocal interactions of plant traits with herbivore damage and pollination across 2 years using the perennial plant Chamerion angustifolium. We measured (1) plant responses to manipulation of damage and pollination in the year of treatment and the subsequent season, (2) damage and pollination responses to manipulation of plant size and flowering traits in the year of treatment, and (3) plant-mediated indirect interactions between herbivores and pollinators. We found that plant traits had little effect on damage and pollination, but damage and pollination affected plant traits in both the treatment year and the subsequent year. We found evidence of indirect effects between leaf herbivores and pollinators in both directions; indirect effects of pollinators on leaf herbivores have not been previously demonstrated. Our results indicate that pollen receipt results in shorter plants with fewer stems but does not change flower number, while leaf herbivory results in taller plants with fewer flowers. Together, herbivory and pollination may contribute to intermediate plant height and plants with fewer stems and flowers in our system.     

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.     

The older plant gets the sun: Age‐related changes in Miscanthus × giganteus phenology

Age‐related changes are usually overlooked in perennial grass research; when they are considered it is usually as a change in plant size (e.g., biomass). Whether other physiological or developmental aspects change as stands age, and how those aspects may impact long‐term stand dynamics, remains unclear. Conventional experimental designs study a single stand over multiple growing seasons and thereby confound age‐related changes with growing season conditions. Here we used a staggered‐start experimental design with three repeated planting years over two growing seasons to isolate growing season effects. We studied changes in Miscanthus × giganteus phenology during its yield‐building stage (first 3 years) and estimated age, growing season and nitrogen (N) effects on development using nonlinear regression parameters. Stand age clearly changed plant growth; faster developmental rates were usually seen in 1‐year‐old stands (young), but because 2‐ and 3‐year‐old stands (mature) emerged 3 months earlier than newly planted stands they produced 30% more stems with 30%–60% more leaves. Nitrogen fertilization modulated some age‐related phenological changes. Fertilized 2‐year‐old stands reached similar stem densities as unfertilized 3‐year‐old stands and had fewer number of senesced leaves like 1‐year‐old stands. In addition, N fertilization had no effect on young M. × giganteus, but extended mature stands’ growing season more than 2 weeks by hastening emergence and delaying senescence. It also delayed flowering regardless of stand age. Our results suggest that, along with changes in size, M. × giganteus stands showed shifts in developmental strategies: young stands emerged later and developed faster, while mature stands grew for longer but more slowly. In temperate regions, where hard frost events are likely to interrupt development in late autumn, rapid early development is critical to plant survival. Nonlinear regression parameter differences proved effective in identifying phenological shifts.     

Effects of a multi-year drought on a drought-adapted shrub, <Emphasis Type="Italic">Artemisia tridentata</Emphasis>

Models of climate change predict more variable precipitation for much of western North America, including more severe multi-year droughts. Droughts are known to increase mortality to trees although less is known about effects on shrubs from arid environments and about effects on reproduction. In this study, we followed a cohort of young sagebrush plants from 2010 to 2016, a period that included a severe drought from 2012 to 2015. Plants experienced little mortality preceding and during the drought. However, in the year following the drought, 14% of individuals died and 33% of branches on living plants died. There was little flowering in the years preceding the drought and flowering increased in each successive year from 2014 to 2016. Plants that produced more flowers in 2015 had more dead branches in 2016. Larger plants had fewer branches that died. Contrary to expectations, afternoon shade was not associated with greater survival or flowering, perhaps because shaded plants were in proximity to large trees which likely competed for water. Plants of the two common chemotypes had similar rates of survival and flowering. Experimental watering during the summer of 2015 did not affect survival and may have increased flowering in 2016. If multi-year droughts become more common in the future, even drought-adapted shrubs may be expected to suffer high rates of mortality.     

Screening a cucumber crop during leaf area development reduces yield

In order to reduce heat energy consumption in greenhouse cucumber production, (transparent) screens may be used also during the day, particularly in the early growth phase when high temperatures are required to achieve rapid leaf area development. However, energy savings must be optimised against light reduction‐induced yield loss. For this reason, two experiments were conducted to quantify the effect on photosynthesis and growth of screening cucumber plants during their early growth phase, and on yield in the following generative phase. Screening with different light transmission coefficients was simulated using shading nets. Shading the plants during the first 5 weeks under Central European winter conditions reduced the leaf area by 0.40% per 1% reduction in photosynthetic active radiation (PAR). Moreover, potential leaf net photosynthesis decreased by 0.46% per 1% PAR reduction. A major impact was that the leaf dry matter content, leaf starch content and leaf sugar content of shaded plants diminished significantly. In the course of the following 2 weeks under full light, the leaf photosynthesis of the plants previously shaded recovered fully and the leaf area index rose to 3.3 m² m^?2, considered sufficient for optimal crop photosynthesis. The yield from plants previously shaded diminished slightly as early as from the first harvest week on. These yield losses increased further over the next few weeks, measuring approximately 0.8 kg m^?2 per 1 mol m^?2 day^?1 PAR reduction in the early growth phase. The effect of PAR on plant growth was proportional when relating the PAR integral over the entire experimental period to the total yield and to the total dry matter production, respectively.     

不同生长阶段遮荫对番茄光合作用、干物质分配与叶N、P、K的影响

对番茄植株做了两种不同程度的遮荫处理,观测了夏季午间遮荫对光合速率,干物质积累量及其在根,茎,叶之间的分配,和叶N,P,K的含量以及经济产量的影响,发现不同时期遮荫影响不同。（1）遮荫增加三个阶段（开花早期,盛花期和开花后期）的气孔导度和胞间CO2浓度,显著降低开花早期中午的净化合速率,但盛花期中度遮荫（40%遮荫）使净光合速率随着时间的增加逐渐上升,在开花后期表现更加明显,平均净光合速率比对照高20%以上,蒸腾速率也增加较多。（2）开花早期和盛花期重度遮荫（如本实验中的75%遮荫）显著降低根,茎的干重,而开花后期中度遮荫的根,茎干重高于对照,但遮荫对叶干重的影响不明显。（3）开花早期和盛花期遮荫不明显影响叶片中N,P,K的含量,但开花后期中度遮荫使N,P,K含量增加,（4）开花早期两种遮荫对果实产量影响较小,但盛花期重度遮荫使产量降低,全部产量中无效部分所占的比例上升,开花后期中度遮荫的总产量和有效产量增加,单果重也增加,这些结果表明,在某些时期中度遮荫可以克服夏天辐射过强,气温过高对番茄的不良影响,对番茄生长,干物质积累和提高产量等有利,在生产上有意义。     

Variation in carbon isotope discrimination in relation to plant performance in a natural population of <Emphasis Type="Italic">Cryptantha flava</Emphasis>

Few studies of phenotypic selection have focused on physiological traits, especially in natural populations. The adaptive significance of plant water-use efficiency, the ratio of photosynthesis to water loss through transpiration, has rarely been examined. In this study, carbon isotopic discrimination, Δ, an integrated measure of water-use efficiency, was repeatedly measured in juveniles and adults in a natural population of the herbaceous desert perennial Cryptantha flava over a 4-year period and examined for plasticity in Δ, consistency between years in values of Δ, and evidence for selection on Δ phenotypes. There was significant concordance in Δ values among the 4 years for adult plants and significant correlations in Δ values measured in different years for juveniles and adults combined. The wettest year of the study, 1998, proved an exception because Δ values that year were not correlated with Δ values in any other year of the study. Consistency in Δ measured on the same plants in different years could indicate genotypic variation and/or consistency in the water status of the microhabitats the plants occupied. Two forms of plasticity in Δ were also evident; mean seasonal values were correlated with precipitation the preceding autumn, and Δ values also declined with plant size, indicating increasing water-use efficiency. Phenotypic selection was evident because in the first year of the study juvenile plants that would survive until year five averaged lower Δ values than did those that failed to survive. During the driest year, 2000, Δ was significantly negatively correlated with adult plant size, measured as the number of leaf rosettes, but the negative relationship between Δ and the number of flowering stalks, a more direct measure of fitness, was not significant. These results suggest that the direction of phenotypic selection on Δ changes as plants grow.     

Demographic consequences of drought in the herbaceous perennial Cryptantha flava: effects of density,associations with shrubs,and plant size

The demographic consequences of a severe drought year were examined for two experimental plantings of the herbaceous desert perennial Cryptantha flava(Boraginaceae) in northeastern Utah, United States. A total of 6680 nutlets were planted individually or in clusters of four both under shrubs and in open microhabitats within two natural populations. Survival, growth, and flowering as a function of density and microhabitat were followed for 7 years, including 1 year when precipitation just before and during the growing season was 74.5% below normal. The design permitted assessment of how intraspecific density and shrub cover affect demographic response to drought. Mortality increased and flowering decreased dramatically during drought but neither varied with density or between shrub and open microhabitats. For plants growing under shrubs, survival (at Site 1) and growth (at Site 2) varied with shrub species. Average aboveground plant size also decreased during drought. Population size hierarchies were rearranged because larger plants lost leaf rosettes while many smaller plants grew. Density and microhabitat affected plant performance in non-drought years but more often at Site 1 than at Site 2. Individuals growing alone often were more likely to flower and/or produced more inflorescences when they did flower than did individuals growing with at least one other plant. However, for 2 years, survival rates at Site 1 were higher for plants growing in clumps than for single individuals. Shrubs also had mixed effects on plant performance. In some years, survival was higher under shrubs, but at Site 1 plants in the open often were more likely to flower and/or produced more inflorescences. Thus despite severe demographic consequences of drought, the study provided no evidence that intraspecific competition, interference by shrubs, or facilitation by shrubs increases under limited soil water.     

In vitro flowering of indica rice (Oryza sativa L. spp. indica)

Nodal explants of rice cultivar Pathumthani 1 (PT1; short-day photoperiod insensitive) were collected, surface-disinfected, and cultured on modified MS medium under in vitro conditions for 90 d. A total of 60% nodal explants generated flowering plantlets (with one inflorescence per cluster). The net photosynthetic rate was greater, and soluble sugars (including glucose, fructose, and sucrose) accumulated to higher levels in the leaves of flowering as compared to non-flowering plants. In contrast, chlorophyll a, chlorophyll b, total chlorophyll, and total carotenoid content were enriched to a greater degree in the leaves of non-flowering as compared to flowering plants. Also, growth performance parameters, including plant height, number of leaves per plant, leaf area, fresh weight, and dry weight of plantlets derived from seedlings were superior to those of plantlets derived from nodal explants. In addition, the protocol proved to successfully induce flowering in KDML 105, a short-day photoperiod-sensitive rice cultivar.     

EFFECTS OF SINGLE AND MULTIPLE EXPOSURES OF FERULIC ACID ON THE VEGETATIVE AND REPRODUCTIVE GROWTH OF PHASEOLUS VULGARIS BBL-290

Phaseolus vulgaris BBL-290 plants were grown in growth chambers in the Southeastern Plant Environment Laboratory and exposed to either single (at seedling, flower, or podfill) or multiple (biweekly or weekly) treatments of ferulic acid (FA). In the first experiment, plants were harvested one week after FA treatment (0, 1.0, 2.0 mM) and at final harvest (56 days old). FA delayed leaf expansion during the seedling and flowering stages. The total plant leaf area and the plant dry weight of plants treated with 1.0 and 2.0 mM FA as seedlings were reduced one week after treatment by 38–48%. The total plant leaf area and the plant dry weight of plants treated at flowering with 2.0 mM FA were reduced by 25% one week after treatment. Treatment with 2.0 mM FA at podfill caused the senescence and abscission of older leaves and reduced total plant leaf area, plant dry weight and mean pod dry weight by 54, 40, and 48%, respectively, one week after treatment. The plants treated at the seedling and flowering stages recovered by final harvest. In a subsequent experiment, FA (0, 0.50, 1.0, 1.5 mM) reduced total plant leaf area at the seedling and flowering stages but not at podfill. The youngest expanding leaves were most sensitive to FA at flowering. The leaf area of these leaves was reduced by 35 and 25%, one and two weeks after treatment, respectively. Their absolute growth rates were reduced from 31 to 56% one week after treatment at flowering. Their relative growth rates were reduced by 50% one week after treatment. Growth rates then recovered within two weeks after treatment. In the final experiment, biweekly exposures of FA (0.25, 0.50, 0.75, 1.0) reduced total plant leaf area but did not affect any other growth parameters. Weekly exposures of FA (0.25, 0.50, 0.75, 1.0) reduced total plant leaf area up to 34%, absolute growth rate up to 58%, leaf number up to 31% and pod number up to 58%. As the frequency of exposure to FA increased, the concentration necessary to affect bean plant growth and development decreased.     

Roles of succession,light, nutrients and disturbance on population vigor and maintenance of the rare plant Solidago shortii (Asteraceae)

Results of field and glasshouse experiments on Solidago shortii, and our observations on this species over many years, were used to construct a conceptual model of the roles of succession, light, soil nutrients and disturbance on population vigor and maintenance of this federal-endangered species. As cover of woody vegetation increased at a population site between 1986 and 1992, number of flowering ramets of S. shortii significantly decreased but number of vegetative ramets remained nearly constant. Adult plants transplanted into a redcedar thicket and those shaded in a glasshouse produced many fewer flowering ramets and capitula per flowering ramet and less biomass and had higher mortality than those in the open. Seedlings/juveniles shaded in a glasshouse had significantly less dry biomass; lower RGR, NAR, leaf area and root/shoot ratio and higher LAR, SLA and LWR than nonshaded ones. In a field site and glasshouse, fertilized plants (NPK) consistently had more flowering ramets and capitula per flowering ramet than nonfertilized ones. Hierarchy of dry weight of plants grown in a glasshouse in soils derived from five types of bedrock was phosphatic limestone > calcareous shale > sandstone > black shale = dolomite. Flowering and biomass production in the field-fertilizer and soil-type experiments were associated closely with levels of P. Number of flowering ramets significantly increased in plants transferred from shaded to nonshaded glasshouse conditions, but no such increase occurred after opening the canopy above plants in a thicket. Both high light and high nutrient levels apparently are necessary to maintain high vigor of S. shortii. In areas subject to invasion by woody plants, periodic high intensity disturbance may be required to prevent population extirpation.     

Is the insect or the plant the driving force in the cinnabar moth — Tansy ragwort system?

Summary The interactions between cinnabar moth, Tyria jacobaeae L. and its food plant, tansy ragwort, Senecio jacobaea L. were studied for 4 to 6 years at 9 sites in North America to assess if the herbivore drove the dynamics of the plants or if the plants determined the dynamics of the insects. Cinnabar moth larval density is not closely related to changes in the size and spacing of tansy ragwort plants although high densities of larvae were associated with a high proportion of rosettes the next year. Fluctuating moth populations live in areas where rosettes are small, closely spaced and numerous compared to flowering stem plants. This situation is also associated with greater larval dispersal which may lead to over exploitation of the food supply. The coefficient of variation of both the size of rosettes and the distance between clumps of plants is associated with the coefficient of variation in moth density. This suggests that the plants may be driving the dynamics of the insect populations. The size of the moth egg batches is correlated to the size of the flowering stem plants in the previous year, indicating an adjustment between moth fecundity and food plant size. The conclusion is that environmental factors such as weather and soil type determine to a large degree the characteristics and variation in the plant populations and this in turn controls the dynamics of the insects. The relation of this situation to the biological control of weeds is discussed.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.     

Physiologic and Yield Responses of Shaded Cotton to the Plant Growth Regulator PGR-IV

The plant growth regulator PGR-IV has been reported to improve the growth, boll retention, and yield of cotton (Gossypium hirsutum L.) under optimum growing conditions. However, little is known about the response of cotton to PGR-IV under low light stress. A 3-year field study was conducted to determine if applying PGR-IV before an 8-day period of shade (63% light reduction) benefitted the growth and yield of shaded cotton. Shading during early squaring did not affect yield. Shading after the first flower stage significantly increased leaf chlorophyll concentration and fruit abscission and decreased the leaf photosynthetic rate, nonstructural carbohydrate concentrations, and lint yield. Foliar application of PGR-IV at 292 mL ha⁻¹ at early squaring and first flower did not improve the leaf photosynthetic rate of shaded cotton. However, shaded plants receiving PGR-IV had higher nonstructural carbohydrate concentrations in the floral buds and significantly lower fruit abscission than the shaded plants without PGR-IV. Applying PGR-IV to the foliage before shading resulted in a numeric increase (6–18%) in lint yield compared with shaded plants without PGR-IV. The decreased fruit abscission from the application of PGR-IV was associated with improved assimilate translocation. The yield enhancement from foliar application of PGR-IV was attributed to increased fruit retention. However, the average boll weight of shaded plants with PGR-IV tended to be lower than that of shaded plants without PGR-IV. Lint percentage was not affected by PGR-IV. Foliar application of PGR-IV appears beneficial for increasing the fruit retention of shaded cotton. Received June 12, 1997; accepted January 19, 1998     

EFFECTS OF AGE AND SIZE ON LIFE HISTORIES AND POPULATION GROWTH OF RHODODENDRON MAXIMUM SHOOTS

Age- and size-specific shoot life histories were studied with population censuses in June 1984 and June 1985 in an evergreen understory shrub. Rhododendron maximum. Most shoots (65%) survived without branching or flowering, and lesser numbers branched (2%), flowered (20%), or died (23%) during the year between censuses. The probabilities of surviving, branching, flowering or dying were both age- and size-dependent. Small, young shoots increased in leaf area. Flowering occurred most prominently in 3- to 6-year old shoots that had exceeded a leaf area of 200 cm², and the rate of flowering increased proportionately with size above this threshold. Branching normally occurred in the year following flowering. The age and size distributions of the population shifted significantly between years, indicating a nonequilibrium population. The survival schedule was Deevey Type I, indicating a high degree of “parental care” of young shoots. Age- and age + size-based demographic models predicted a rapid decline of the shoot population over a decade, while a size-based model predicted a much slower decline in shoot numbers. A sensitivity analysis of the models showed that overall shoot population growth was positively influenced by branching shoots and shoots that added leaf area, and negatively influenced by shoots that lost leaf area, died, or flowered. The role of shoot life histories in determining individual plant fitness and ecological dominance is discussed.