Resource availability and growth responses to defoliation in seedlings of three early-successional,tropical, woody species

A growth experiment was conducted using seedlings of three early-successional, tropical, woody species:Dillenia suffruticosa (Dilleniaceae),Macaranga heynei (Euphorbiaceae) andTrema tomentosa (Ulmaceae). These species are characteristic of different positions along a soil fertility gradient in Singapore, withD. suffruticosa being the least andT. tomentosa the most demanding of high nutrient availability. The seedlings were grown in vermiculite at either low or high nutrient availabilities supplied by watering with different concentrations of a commercial plant food. Half the seedlings were subjected to a 50% defoliation at the start of the experiment by cutting off the distal half of each leaf. After 9 weeks the plants were harvested.Macaranga heynei andT. tomentosa seedlings showed no significant difference in parameters of growth such as total dry weight and total leaf area between the defoliated and control seedlings. The seedlings compensated completely for the loss of leaf area.Dillenia suffruticosa did show significant reductions in growth in some parameters due to defoliation, and these were more pronounced under the high nutrient treatment. These findings support the hypothesis that plants characteristic of resource-rich sites can readily recover from herbivory through fast growth, probably associated with a rapid turnover of leaves, whereas species of resource-poor habitats cannot easily replace losses due to herbivory and are adversely effected by defoliation.     

Density- and growth stage-dependent responses to defoliation in two rhizomatous grasses

Summary Responses to defoliation were studied in two tallgrass prairie perennials (Andropogon gerardii and Panicum virgatum) established from seed at three densities. P. virgatum was also grown from transplanted rhizomes of established clones. Plants of both species displayed a continuum of responses to defoliation, from large reductions in biomass, tillering and seed production to significant increases in one or more performance measures. In crowded populations, defoliation shifted plants into subordinate positions within the competitive hierarchy. Plants competing intraspecifically and those that were initially small suffered more from defoliation than either plants grown at low density or those that were larger than their neighbors. At the highest plant density, the effects of defoliation or initial plant size were overshadowed by the effects of crowding. When defoliated and grown at similar densities, P. virgatum and A. gerardii grown from seed showed large reductions in biomass, seed production, and new rhizome production, but established P. virgatum ramets grown from rhizomes showed increases in these performance measures. Thus, herbivory may be particularly detrimental to P. virgatum during juvenile stages before perennating organs have developed. Overcompensation of P. virgatum clones in response to defoliation only occurred if all ramets within the clone were defoliated. In clones containing both defoliated and undamaged ramets, there were no differences in their performance, suggesting that genets are capable of integrating the effects of differential defoliation among shoots. Defoliated P. virgatum clones allocated a smaller fraction of their total biomass to new rhizomes, indicating that the short-term regrowth response following defoliation may incur a longer-term cost associated with gradual reduction in biomass of the perennating organs and reduced genet success.     

Comparative responses of the Savanna grasses Cenchrus ciliaris and Themeda triandra to defoliation

Summary Two perennial tussock grasses of savannas were compared in a glasshouse study to determine why they differed in their ability to withstand frequent, heavy grazing; Cenchrus ciliaris is tolerant and Themeda triandra is intolerant of heavy grazing. Frequent defoliation at weekly intervals for six weeks reduced shoot biomass production over a subsequent 42 day regrowth period compared with previously undefoliated plants (infrequent) in T. triandra, but not in C. ciliaris. Leaf area of T. triandra expanded rapidly following defoliation but high initial relative growth rates of shoots were not sustained after 14 days of regrowth because of reducing light utilising efficiency of leaves. Frequently defoliated plants were slower in rate of leaf area expansion and this was associated with reduced photosynthetic capacity of newly formed leaves, lower allocation of photosynthate to leaves but not lower tiller numbers. T. triandra appears well adapted to a regime where defoliation is sufficiently infrequent to allow carbon to be fixed to replace that used in initial leaf area expansion. In contrast, C. ciliaris is better adapted to frequent defoliation than is T. triandra, because horizontally orientated nodal tillers are produced below the defoliation level. This morphological adaptation resulted in a 10-fold higher leaf area remaining after defoliation compared with similarly defoliated T. triandra, which together with the maintenance of moderate levels of light utilising efficiency, contributed to the higher leaf area and shoot weight throughout the regrowth period.     

Ecophysiological responses to light availability in three Blechnum species (Pteridophyta, Blechnaceae) of different ecological breadth

In Chilean evergreen temperate forest, fern species of the genus Blechnum occur in diverse microhabitats ranging from large gaps to heavily shaded understoreys. We hypothesised that differences in the ecological breadth of three co-occurring Blechnum species would be associated with differences in magnitude of ecophysiological responses to light availability. We quantified the field distribution of each species in relation to diffuse light availability (% canopy openness), and measured in situ variation in photosynthetic capacity (A), dark respiration (R (d)) and specific leaf area (SLA) across the light gradient. The response of SLA of each species was also evaluated in a common garden in two light conditions (understorey and forest edge). The three Blechnum species differed significantly in the range of light environments occupied (breadth: B. chilense > B. hastatum > B. mochaenum). Despite significant interspecific differences in average A and R (d), the response of these traits to light availability did not differ among species. However, there was significant interspecific variation in both the mean value and the plasticity of SLA to light availability, the species with least ecological breadth (B. mochaenum) showing a flatter reaction norm (lower response) than its two congeners. This pattern was also found in the common garden experiment. The adjustment of leaf morphology (SLA) to light availability appears to be an important mechanism of acclimation in these Blechnum species. The narrow range of light environments occupied by B. mochaenum may be at least partly attributable to its inability to display phenotypic plasticity in SLA to changes in light availability.     

Effect of defoliation on the arbuscular mycorrhizas of three perennial pasture and rangeland grasses

The effect of different frequencies of defoliation on arbuscular mycorrhizal fungal (AMF) colonization and external hyphae production of three perennial grass species growing in pot culture in a non-sterile soil was investigated. Roots were assessed by acid fuschin staining and succinate dehydrogenase activity to obtain measurements of total and metabolically active AMF colonization. The grass species, Digitaria eriantha, Lolium perenne and Themeda triandra are of similar bunch morphology and responded to defoliation with massive root death. In Themeda defoliation was also associated with a decline in leaf growth rate, phosphorus accumulation in new leaf tissue, AMF colonization and external hyphae densities. In Digitaria and Lolium, AMF colonization declined but external hyphal densities were unaffected by defoliation frequency. In these two species phosphorus accumulation and leaf regrowth rates were also unchanged by defoliation. Only in Lolium did defoliation result in slightly more inactive AMF colonization. The results suggest that Lolium and Digitaria which are pasture species are better able to compensate for root loss following fairly frequent defoliation by maintaining an external AMF hyphal network. Themeda, a rangeland grass, which is more intolerant of grazing, has a lower capacity for sustaining its hyphal network when defoliated. Grazing is therefore likely to affect community dynamics because of variable effects of defoliation frequency on the mycorrhizal symbiosis of different plant species.     

Ecophysiological responses to light availability in three Blechnum species (Pteridophyta, Blechnaceae) of different ecological breadth

In Chilean evergreen temperate forest, fern species of the genus Blechnum occur in diverse microhabitats ranging from large gaps to heavily shaded understoreys. We hypothesised that differences in the ecological breadth of three co-occurring Blechnum species would be associated with differences in magnitude of ecophysiological responses to light availability. We quantified the field distribution of each species in relation to diffuse light availability (% canopy openness), and measured in situ variation in photosynthetic capacity (A), dark respiration (R _d) and specific leaf area (SLA) across the light gradient. The response of SLA of each species was also evaluated in a common garden in two light conditions (understorey and forest edge). The three Blechnum species differed significantly in the range of light environments occupied (breadth: B. chilense > B. hastatum > B. mochaenum). Despite significant interspecific differences in average A and R _d, the response of these traits to light availability did not differ among species. However, there was significant interspecific variation in both the mean value and the plasticity of SLA to light availability, the species with least ecological breadth (B. mochaenum) showing a flatter reaction norm (lower response) than its two congeners. This pattern was also found in the common garden experiment. The adjustment of leaf morphology (SLA) to light availability appears to be an important mechanism of acclimation in these Blechnum species. The narrow range of light environments occupied by B. mochaenum may be at least partly attributable to its inability to display phenotypic plasticity in SLA to changes in light availability.     

Differential effects of interspecific interactions and water availability on survival, growth and fecundity of three congeneric grassland herbs

Fitness of individual plants and of populations depends on the rates of survival, growth and fecundity. This study tested whether vital rates were differentially affected by biotic interactions and water availability. The effects of manipulations of above-ground competition (through clipping) and water availability (through water addition) on the vital rates of seedlings of three species (Viola elatior Fries, Viola pumila Chaix and Viola stagnina Kit.) were analysed in dry, mesic and wet grasslands. Water addition and grassland type had the largest effects on survival (accounting for 41 and 24% of total variation, respectively) across species. Height growth rate was positively affected by grassland type (19%) and water addition (12%) and varied among species (8%), while leaf accumulation rates and reproduction were affected by grassland type and clipping. The data suggested facilitative effects of the canopy on seedling survival in the dry grassland. This study presents evidence that environmental conditions and biotic interactions may have differential effects on seedling survival, growth and reproduction. The findings highlight the complex interplay between spatial and temporal environmental variation and biotic interactions in structuring plant communities.     

Semiarid grassland responses to short-term variation in water availability

Standing crop and species composition in semiarid grassland are linked to long-term patterns of water availability, but grasslands are characterized by large single-season variability in rainfall. We tested whether a single season of altered water availability influenced the proportions of grasses and shrubs in a semiarid grassland near the northern edge of the North American Great Plains. We studied stands of the clonal shrub snowberry (Symphoricarpos occidentalis) and adjacent grassland dominated by the native grasses Stipa spartea and Bouteloua gracilis. Rain was excluded and water supplied in amounts corresponding to years of low, medium, and high rainfall, producing a 2 − 4-fold range in monthly precipitation among water supply treatments. There were ten replicate plots of each water treatment in both snowberry stands and grassland. Grass standing crop increased significantly with water availability in grassland but not inside snowberry stands. Total standing crop and shrub stem density increased significantly with water supply, averaged across both communities. In contrast, water had no effect on shrub standing crop or light penetration. In summary, our finding that water has significant effects on a subset of components of grassland vegetation is consistent with long-term, correlational studies, but we also found that a single season of altered water supply had no effect on other important aspects of the ecosystem. This revised version was published online in June 2006 with corrections to the Cover Date.     

Modeling the influence of differential sectoriality on the photosynthetic responses of understory saplings to patchy light and water availability

Exploitation of patchy light is a key determinant of plant performance in the forest understory. While many adaptive traits are known, the role of stem vasculature in understory photosynthesis is not established. Sectoriality—the degree of vascular constraint to long distance transport—has been hypothesized to limit growth in heterogeneous light. We simulated the photosynthetic potential of sectored and integrated plants in patchy light, as a function of soil water potential (patchy or uniform). We used hydraulic parameters typical of temperate woody species in an Ohm’s law model including a tangential resistance parameter, and simulated cavitation by varying axial resistance of leaves, leaves and roots, or the whole plant. Our results suggest that differential sectoriality will not affect photosynthesis when water is plentiful, but can constrain stomatal conductance at more negative soil water potentials, especially when only a small portion of the crown receives light. This effect is strongest just below the turgor loss point, and depends on axial resistance and soil water heterogeneity. Increased resistance in high light leaves decreases photosynthesis regardless of sectoriality. However, when resistance is increased for leaves and roots or the whole plant, photosynthesis decreases more for sectored than for integrated plants. Moreover, the simulations suggest that sectoriality can further depress photosynthesis when water availability is asymmetrical. These results might explain why integrated species, such as Betula lenta, B. alleghaniensis, and Acer saccharum thrive in the forest understory and grow rapidly into canopy gaps, while sectored species, such as Quercus rubra, do not.     

Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

Morphological and physiological responses of two coffee progenies to soil water availability

Drought is a major environmental constraint affecting growth and production of coffee. The effects of water supply on growth, biomass allocation, water relations, and gas exchange in two coffee progenies representing drought-tolerant (Siriema) and drought-sensitive (Catucaí) genotypes were compared. They were grown in 12-L pots until 4-months old, when they were submitted to two watering treatments for 60 d: plants receiving either 100% transpired water (control plants) or a fraction (about 40%) of the amount of water transpired by control plants (drought-stressed plants). Under control conditions, Siriema grew faster than Catucaí. Regardless of the watering regimes and progenies, relative growth rate (RGR) was positively correlated both with net assimilation rate (NAR) and long-term water-use efficiency (WUE), but not with differences in biomass allocation. Both progenies responded to drought stress through (i) similar decreases in both RGR and NAR with marginal, if any, changes in allocation; (ii) decreases in leaf water potential, which occurred to a greater extent in Catucaí than in Siriema, even though they have showed similar abilities to adjust osmotically and elastically; (iii) similar reductions in net photosynthesis due mainly to nonstomatal factors; and (iv) decreases in transpiration rate coupled with increased long-term WUE. However, the lower transpiration rate and the higher long-term WUE as found in Siriema relative to Catucaí under control conditions persisted under drought conditions. Overall, the major differences between these progenies were largely associated with differences in plant water use, which was likely related to the improved water status of Siriema. The possible implications of selecting coffee genotypes for high WUE are discussed.     

Leaf optical responses to light and soil nutrient availability in temperate deciduous trees

Leaf optical parameters influence light availability at the cellular, leaf, and canopy scale of integration. While recent studies have focused on leaf optical responses to acute plant stress, the effects of changes in plant resources on leaf optics remain poorly characterized. We examined leaf optical and anatomical responses of five temperate deciduous tree species to moderate changes in nutrient and light availability. Spectral reflectance in the visible waveband generally increased at high light, but decreased with increased nutrient availability. Patterns of both spectral reflectance and absorptance were primarily determined by chlorophyll concentration although carotenoid concentration was also influential. While most anatomical features did not explain residual variation in reflectance, cuticle thickness was significantly related to reflectance at complementary angles compared to the angle of incidence. Absorptance did not change with light environment; however, absorption efficiency per unit biomass increased by approximately 40% under low light, due to reduced leaf mass per area. We conclude that changes in resource availability differentially influence leaf optical properties and that such changes are driven primarily by changes in pigment concentrations. The magnitude of leaf optical responses to moderate changes in resource availability was comparable to those of acute stress responses and varied among species.     

Differential physiological responses to prey availability by the great egret and white Ibis

In long-lived species, the balance between the benefits of reproduction and the costs from reduced survival or productivity is particularly challenging in dynamic environments like wetlands, where food levels vary greatly year to year. Some wetland species exhibit changes in reproductive strategies in response to food availability but whether physiological responses function in a similar manner is unclear. We compared the pre-breeding physiological responses (fecal corticosterone [FCORT], heat shock protein 60 [HSP60], and mass) of 2 species of wading birds with contrasting foraging strategies (great egret [Ardea alba], an exploiter, and white ibis [Eudocimus albus], a searcher) during years with contrasting levels of prey availability. Both species were in good physiological condition, with low levels of HSP60 and FCORT, during a year with high prey availability (2006). In a contrasting year with lesser prey availability (2007), HSP60 and FCORT concentrations indicated that ibis physiological condition was reduced, whereas egrets showed little change. Egrets and male ibis increased body mass, whereas female ibis decreased mass, in the year with low prey availability. Although poorly understood, we hypothesize that the differential response between female ibis and the others is associated with differential investment strategies based on long-term costs of reproduction. Model results identified prey availability and the 2-week water recession rate as the primary habitat variables that were associated with the physiological condition of white ibises, whereas great egret physiological condition was influenced mostly by 2-week water recession rate. Our results support the hypothesis that prey availability and hydrological factors play crucial roles in regulating populations of wading birds in the Florida Everglades. The results of this study show a more complete pathway by which hydrologic patterns affect wading birds, and it suggests that ibis are more sensitive to habitat conditions than are egrets. This information can be used to refine species models designed to evaluate water management scenarios and will improve our ability to manage and restore wetland ecosystems © 2012 The Wildlife Society.     

Adaptation of selected trees and grasses to low availability of phosphorus

High-nutrient-adapted and low-nutrient-adapted species of New Zealand tussock grasses (Chionochloa), barley (Hordeum), and several taiga trees were grown at three rates of phosphorus supply. Low-nutrient-adapted species in each group of species had similar (grasses) or lower (trees) capacities for phosphate absorption, were less efficient in producing biomass (i.e. had higher nutrient concentrations), and grew more slowly than high-nutrient-adapted species. I conclude that the major adaptation to low nutrient availability in each of these comparisons is a slow growth rate that reduces the annual nutrient requirement.     

Root traits associated with nutrient exploitation following defoliation in three coexisting perennial grasses in a semi-arid savanna

Experiments were conducted to evaluate root traits associated with nutrient exploitation following defoliation in three coexisting perennial grasses in a semi‐arid savanna. Root length density was determined within soil cores directly beneath plants, nitrogen uptake was evaluated by excised‐root assay with (¹⁵NH₄)₂SO₄, and mycorrhizal root colonization was estimated by observation of root segments. Root length density was lowest for Bouteloua curtipendula, intermediate for Eriochloa sericea, and highest for Aristida purpurea indicating that root length density was a more important trait for the mid‐seral than the late‐seral species. Rates of ¹⁵N uptake were greatest in the least grazing tolerant late‐seral species, E. sericea, intermediate in the mid‐seral species, A. purpurea, and lowest in the most grazing tolerant late‐seral species, B. curtipendula. Two successive defoliations reduced ¹⁵N uptake 60% in the late‐seral species with the greatest uptake rate (E. sericea), but not in species with lowest uptake rates (B. curtipendula). Root length colonization was consistently high (33–61%) in all three species suggesting that these C₄ perennial grasses may function as obligate mycotrophs. Contrasting responses among the two late‐seral species indicate that the least grazing tolerant species, E. sericea, appears best adapted for nutrient exploitation while the most grazing tolerant species, B. curtipendula, appears best adapted for efficient nutrient retention. Contrasting responses of nitrogen uptake to short‐term defoliation parallel the population responses of these two coexisting late‐seral species to long‐term herbivory. These data indicate that herbivory may shift interspecific competitive interactions by mediating nutrient exploitation and that a trade‐off may exist between nutrient exploitation and herbivory tolerance in these species.     

Proteomic profiling of aphid Macrosiphum euphorbiae responses to host-plant-mediated stress induced by defoliation and water deficit

Abiotic and biotic host-plant stress, such as desiccation and herbivory, may strongly affect sap-sucking insects such as aphids via changes in plant chemicals of insect nutritional or plant defensive value. Here, we examined (i) water deprivation and (ii) defoliation by the beetle Leptinotarsa decemlineata as stresses indirectly affecting the aphid Macrosiphum euphorbiae via its host plant Solanum tuberosum. For plant-induced stress, aphids were reared on healthy vs. continuously stressed potato for 14 days (no watering; defoliation maintained at approximately 40%). Aphid performance under stress was correlated with metabolic responses monitored by profiling of the aphid proteome. M. euphorbiae was strongly affected by water stress, as adult survival, total aphid number and biomass were reduced by 67%, 64%, and 79%, respectively. Aphids performed normally on defoliated potato, indicating that they were unaffected or able to compensate any stress induced by plant defoliation. Stressed aphid proteomes revealed 419-453 protein spots, including 27 that were modulated specifically or jointly under each kind of host-plant stress. Reduced aphid fitness on water-stressed plants mostly correlated with modulation of proteins involved in energy metabolism, apparently to conserve energy in order to prioritize survival. Despite normal performance, several aphid proteins that are known to be implicated in cell communication were modulated on defoliated plants, possibly suggesting modified aphid behaviour. The GroEL protein (or symbionin) of the endosymbiont Buchnera aphidicola was predominant under all conditions in M. euphorbiae. Its expression level was not significantly affected by aphid host-plant stresses, which is consistent with the high priority of symbiosis in stressed aphids.     

Sex-specific physiological, allocation and growth responses to water availability in the subdioecious plant Honckenya peploides

The gender of dimorphic plant species is often affected by ecophysiological variables. Differences have been interpreted as a response of the sexes to meet specific resource demands associated with reproduction. This study investigated whether sex‐specific variations in ecophysiological traits in response to water availability determine the performance of each sex in different habitats, and therefore promote extreme spatial segregation of the sexes in the subdioecious plant, Honckenya peploides. Twenty‐seven plants of each sex were individually potted in dune sand and assigned randomly to one of three water treatments. Well‐watered plants were watered daily to field capacity, whereas plants in the moderate and high‐water stress treatments received 40% and 20%, respectively, of the water given to well‐watered plants. Photochemical efficiency, leaf spectral properties and components of relative growth rate (leaf area ratio and net assimilation rate) were measured. Photochemical efficiencies integrated over time were higher in male than in female plants. Water deficit decreased maximum quantum yield in female plants more rapidly than in male plants, but female plants (unlike male plants) had recovered to initial values by the end of the experiment. Maximum quantum yield in male plants was more affected by water stress than in female plants, indicating that male plants were more susceptible to photoinhibition. The two sexes did not differ in growth rate, but male plants invested a higher proportion of their biomass in leaves, had a higher leaf area per unit biomass and lower net assimilation rate relative to female plants. Female plants had a higher water content and succulence than male plants. Differences in stomatal density between the sexes depended on water availability. The results suggest that the two sexes of H. peploides have different strategies for coping with water stress. The study also provides evidence of sex differences in allocation traits. We conclude that between‐sex differences in ecophysiological and allocation traits may contribute to explain habitat‐related between‐sex differences in performance and, therefore, the spatial segregation of the sexes.     

Electrical signals in avocado trees: Responses to light and water availability conditions

Plant responses to environmental changes are associated with electrical excitability and signaling; automatic and continuous measurements of electrical potential differences (ΔEP) between plant tissues can be effectively used to study information transport mechanisms and physiological responses that result from external stimuli on plants. The generation and conduction of electrochemical impulses within plant different tissues and organs, resulting from abiotic and biotic changes in environmental conditions is reported. In this work, electrical potential differences are monitored continuously using Ag/AgCl microelectrodes, inserted 5 mm deep into sapwood at two positions in the trunks of several Avocado trees. Electrodes are referenced to a non polarisable Ag/AgCl microelectrode installed 20 cm deep in the soil. Systematic patterns of ΔEP during absolute darkness, day-night cycles and different conditions of soil water availability are discussed as alternative tools to assess early plant stress conditions.Key words: plant electrical potential, light and dark cycles, water availability, plant signaling, avocado trees, plant sensors, irrigation automation