The potential for and constraints on the evolution of compensatory ability in Asclepias syriaca

To investigate the potential for and constraints on the evolution of compensatory ability, we performed a greenhouse experiment using Asclepias syriaca in which foliar damage and soil nutrient concentration were manipulated. Under low nutrient conditions, significant genetic variation was detected for allocation patterns and for compensatory ability. Furthermore, resource allocation to storage was positively, genetically correlated both with compensatory ability and biomass when damaged, the last two being positively, genetically correlated with each other. Thus, in the low nutrient environment, compensatory ability via resource allocation to storage provided greater biomass when damaged. A negative genetic correlation between compensatory ability and plant biomass when undamaged suggests that this mechanism entailed an allocation cost, which would constrain the evolution of greater compensatory ability when nutrients are limited. Under high nutrient conditions, neither compensatory ability nor allocation patterns predicted biomass when damaged, even though genetic variation in compensatory ability existed. Instead, plant biomass when undamaged predicted biomass when damaged. The differences in outcomes between the two nutrient treatments highlight the importance of considering the possible range of environmental conditions that a genotype may experience. Furthermore, traits that conferred compensatory ability did not necessarily contribute to biomass when damaged, demonstrating that it is critical to examine both compensatory ability and biomass when damaged to determine whether selection by herbivores can favor the evolution of increased compensation. Received: 2 April 1999 / Accepted: 21 September 1999     

EXPERIMENTAL EVOLUTION OF RESISTANCE IN BRASSICA RAPA: CORRELATED RESPONSE OF TOLERANCE IN LINES SELECTED FOR GLUCOSINOLATE CONTENT

The evolutionary response of plant populations to selection for increased defense may be constrained by costs of defense. The purpose of this study was to investigate such constraints on the evolution of defense due to a cost of defense manifested as a trade-off between defense and tolerance. Variation in the response to artificial damage (tolerance) among lines of Brassica rapa that had been artificially selected for foliar glucosinolate content (defense) was examined. Leaf area was removed from replicates of three selection lines (high glucosinolates, control, and low glucosinolates) at three damage levels (0%, 20%, and 60% damage). An external cost of defense would result in a statistically significant selection line by damage treatment interaction, with those selected for high defense expressing less tolerance than those selected for low defense. Damage treatment had a significant overall effect on estimated total fitness, with fitness declining with increasing damage level. Further, selection line also had a significant overall effect on estimated total fitness, with low-defense selection lines having higher fitness compared to both control and high-defense selection lines. More importantly, a cost of defense in terms of tolerance was demonstrated by a significant selection line-by-damage treatment interaction. This interaction was in the direction to demonstrate a genetic trade-off between defense and tolerance, with low-defense selection lines decreasing estimated total fitness in response to damage less than both control and high-defense selection lines. Variation in tolerance among selection lines was due to the greater ability of low-defense lines to maintain fruit and seed production despite the presence of damage. In terms of tolerance, this cost of glucosinolate production in B. rapa could constrain the evolution of increased defense and, in so doing, maintain individuals within the population that are poorly defended yet tolerant.     

Intraspecific variation in the response of Themeda triandra to defoliation: the effect of time of recovery and growth rates on compensatory growth

Summary The response to a single defoliation was studied on three clones of Themeda triandra collected in the short, mid, and tall grassland regions of the Serengeti National Park (Tanzania). These sites represent a gradient of decreasing grazing intensity. Growth, allocation pattern, and several morphometric traits were monitored during an 80-day period. Clipped plants of the short and medium clones fully compensated for the reduction of biomass, while plants of the tall clone showed overcompensation. During the first two weeks after clipping, clipped plants showed lower relative growth rates than unclipped ones, whereas the opposite was observed later on. Clipped plants compensated for the removal of leaf area by producing new leaves with lower specific weights and higher nitrogen content. They also produced more, smaller tillers. Although clipped plants mobilized nonstructural carbohydrates from roots and crowns, this did not account for a significant amount of growth. Relative growth rates of unclipped plants of the short clone were higher. The relative growth rate of the short clone diminished less after clipping, but also exhibited the lowest increase later. The tall clone was the most negatively affected early, but showed the highest compensation later. Compared to the other clones, the short ecotype showed many of the characteristics that defoliation induced in each individual of any clone: higher allocation to leaf area production, higher relative growth rate, higher number but smaller size of tillers, and lower leaf specific weights.     

Scales and costs of habitat selection in heterogeneous landscapes

Summary Two scales of habitat selection are likely to influence patterns of animal density in heterogeneous landscapes. At one scale, habitat selection is determined by the differential use of foraging locations within a home range. At a larger scale, habitat selection is determined by dispersal and the ability to relocate the home range. The limits of both scales must be known for accurate assessments of habitat selection and its role in effecting spatial patterns in abundance. Isodars, which specify the relationships between population density in two habitats such that the expected reproductive success of an individual is the same in both, allow us to distinguish the two scales of habitat selection because each scale has different costs. In a two-habitat environment, the cost of rejecting one of the habitats within a home range can be expressed as a devaluation of the other, because, for example, fine-grained foragers must travel through both. At the dispersal scale, the cost of accepting a new home range in a different habitat has the opposite effect of inflating the value of the original habitat to compensate for lost evolutionary potential associated with relocating the home range. These costs produce isodars at the foraging scale with a lower intercept and slope than those at the dispersal scale.Empirical data on deer mice occupying prairie and badland habitats in southern Alberta confirm the ability of isodar analysis to differentiate between foraging and dispersal scales. The data suggest a foraging range of approximately 60 m, and an effective dispersal distance near 140 m. The relatively short dispersal distance implies that recent theories may have over-emphasized the role of habitat selection on local population dynamics. But the exchange of individuals between habitats sharing irregular borders may be substantial. Dispersal distance may thus give a false impression of the inability of habitat selection to help regulate population density.     

Effect of stress and time for recovery on the amount of compensatory growth after grazing

Summary We tested the hypothesis that the amount of compensatory growth after defoliation is affected by the level of stress at which plants grow when defoliated and by the length of time for recovery. Growth response to defoliation went from partial compensation when plants were growing at high relative growth rates (RGR) to overcompensation when plants were more stressed and growing at low RGR. Defoliation released plants from the limitation imposed by the accumulation of old and dead tissue and this release overrode the negative effect of biomass loss. Compensatory growth resulted from a higher RGR aboveground that was not associated with a reduction in RGR belowground. Time available for recovery had a major impact on the outcome of defoliation. With a short time for recovery, RGR was decreased by defoliation because an immediate increase in net assimilation rate was overridden by a reduction in the ratio of leaf area to plant weight. After defoliation, this ratio increased quickly due to a larger allocation to leaf growth and lower leaf specific weights, resulting in higher RGR. We conclude that the compensatory response to grazing depends on the type and level of stress limiting growth. Allocation and physiological responses to stress may positively or negatively affect the response to grazing and, simultaneously, grazing may alleviate or aggravate the effects of different types of stress.     

Components of relative growth rate and sensitivity to nitrogen availability in annual and perennial species of Bromus

Summary Two grass species, the annual Bromus sterilis and the perennial Bromus erectus, were grown from seeds for 28 days in a hydroponic culture system at 1 and 100 M NO₃ ^- in the nutrient solution. At 100 M NO₃ ^-, the relative growth rate (RGR) of the perennial was 30% lower than that of the annual. This was only the consequence of the higher specific mass of its leaves, since its leaf mass ratio was higher than that of the annual and the unit leaf rates (ULR), calculated on an area basis, were similar for the two species. At 1 M, the RGR of the annual was 50% lower than at 100 M, while that of the perennial was not significantly lower. This was due mainly to a lower ULR for the annual. while for the perennial ULR was the same in both treatments. These differences between the two species were all the more striking in that the differences in total nitrogen concentrations and nitrate reductase activities between the two treatments were very similar for both species. These different responses together with differences in the nitrogen productivity of the two species suggest that the level of nutrient availability may play an important role in the distribution of these Bromus species in natural habitats. Scope: Components of growth and response to nitrate availability in annual and perennial grasses     

The relationship between the relative growth rate and nitrogen economy of alpine and lowland Poa species

This study investigates the nitrogen economy of six altitudinally contrasting Poa species which differ in their relative growth rate (R). Two alpine (Poa fawcettiae and P. costiniana), one sub-alpine (P. alpina)and three temperate lowland species (P. pratensis, P. campressa and P. trivialis) were grown hydroponically under identical conditions in a growth room. The low R exhibited by the alpine species was associated with lower plant organic nitrogen concentration (n_p) and lower nitrogen productivity (Π_p, amount of biomass accumulation per mol organic nitrogen and time). The differences in Π_p between the alpine and lowland species did not appear to be due to differences in the carbon concentration or the proportion of total plant organic nitrogen allocated to the leaves, stems or roots. Variations in Π_P were also not due to variations in photosynthetic nitrogen use efficiency (Ψ_N, the rate of photosynthesis per unit organic leaf nitrogen) or shoot or root respiration rates per unit organic nitrogen (Λ_SH and Λ_R, respectively) per se. Rather, the lower Λ_p in the alpine species was probably due to a combination of small variations in several of the parameters (e.g. slightly lower Ψ_N, slightly higher Λ_SH and Λ_R, and slightly higher proportions of total plant organic nitrogen allocated to the roots). The alpine species exhibited lower organic acid and mineral concentrations. However, no differences in whole-plant construction costs (grams of glucose needed to synthesize one gram of biomass) were observed between She alpine and lowland Poa species. The lack of sub-stantial differences in Ψ_N between the alpine and lowland species contrasts with the large differences in Ψ_N between slow- and fast-growing lowland species that have been reported in the literature. The reasons for the unusually high Ψ_N values exhibited by the alpine Poa species are discussed.     

Dynamics of energy and nutrient concentration and construction cost in a native and two alien C4 grasses from two neotropical savannas

In Venezuela, the alien grasses Melinis minutiflora Beauv. and Hyparrhenia rufa (Nees.) Stapf tend to displace the native savanna plant community dominated by Trachypogon plumosus (Humb. and Bonpl.) Nees. This occurs in either relatively wetter and fertile highland savannas or in drier and less fertile lowland savannas. Although the native and aliens are perennial C₄ grasses, higher net assimilation leaf biomass per plant and germination rate of the latter are some causes for their higher growth rates and for their competitive success. The objective of this study is to compare seasonal tissue energy, N, P and K concentrations and the calculated construction costs (CC) between the native grass and either one of the alien grasses from lowland and highland savannas. We predict that, in order to out-compete native plants, alien grasses should be more efficient in resource use as evidenced by lower tissue energy and nutrient concentrations and CC.Tissue energy and nutrient concentration were measured throughout the year and compared between M. minutiflora and the co-occurring local population of T. plumosus in a highland savanna and between H. rufa and its neighbor local population of T. plumosus in a lowland savanna. CC was calculated from energy, N and ash concentrations considering ammonium as the sole N source. Differences between co-occurring species, T. plumosus populations, seasons, and organs were analyzed with ANOVA.Highland and lowland grasses differed in concentration and allocation of energy and nutrients whereas the differences between alien and native grasses were specific for each pair considered. Highland grasses had higher energy, N, P and CC than lowland grasses. These variables were always lowest in the culms. In the more stressed lowland site, tissue energy and nutrient concentrations decreased significantly during the dry season except in the roots of both grasses which had the highest energy and nutrients concentrations during the drought. This seasonal response was more marked in the local lowland population of T. plumosus in which maximum CC alternated seasonally between leaves and roots. Energy and nutrient concentrations and CC were the lowest in H. rufa. In the lowland savannas, the higher efficiency of resource use in the invader grass contributes to its higher competitive success through increased growth rate. In the highlands, overall tissue energy concentration and CC, but not N nor P concentration, were lower in the fast growing M. minutiflora but seasonal differences were lacking. The higher leaf CC in T. plumosus can be attributed to the higher proportion of sclerenchyma tissue which is more expensive to construct. Considering CC, both fast growing alien grasses are more efficient in resource use than the co-occurring native grass. However, the role of CC explaining the competitive success of the former, through higher growth rates, is more evident in the more stressful environment of the lowland savanna.