Coping with herbivory: Photosynthetic capacity and resource allocation in two semiarid Agropyron bunchgrasses

Summary Agropyron desertorum, a grazing-tolerant bunchgrass introduced to the western U.S. from Eurasia, and Agropyron spicatum, a grazing-sensitive bunchgrass native to North America, were examined in the field for photosynthetic capacity, growth, resource allocation, and tiller dynamics. These observations allowed identification of physiological characteristics that may contribute to grazing tolerance in semiarid environments. A uniform matrix of sagebrush, Artemisia tridentata, provided an ecologically relevant competitive environment for both bunch-grass species. Physiological activity, growth, and allocation were also followed during recovery from a severe defoliation treatment and were correlated with tiller dynamics.Potential photosynthetic carbon uptake of both species was dominated by stems and leaf sheaths during June, when maximum uptake rates occurred. For both species, water use efficiency of stems and sheaths was similar to that of leaf blades, but nitrogen investment per photosynthetic surface area was less than in blades. In addition, soluble carbohydrates in stems and sheaths of both species constituted the major labile carbon pools in control plants. Contrary to current theory, these findings suggest that culms from which leaf blades have been removed should be of considerable value to defoliated bunchgrasses, and in the case of partial defoliation could provide important supplies of organic nutrients for regrowth. These interpretations, based on total pool sizes, differ markedly from previous interpretations based on carbohydrate concentrations alone, which suggested that crowns contain large carbohydrate reserves. In this study, crowns of both species contained a minor component of the total plant carbohydrate pool.Following defoliation, A. desertorum plants rapidly reestablished a canopy with 3 to 5 times the photosynthetic surface of A. spicatum plants. This difference was primarily due to the greater number of quickly growing new tillers produced following defoliation. Agropyron spicatum produced few new tillers following defoliation despite adequate moisture, and carbohydrate pools that were equivalent to those in A. desertorum.Leaf blades of regrowing tillers had higher photosynthetic capacity than blades on unclipped plants of both species, but the relative increase, considered on a unit mass, area, or nitrogen basis, was greater for A. desertorum than for A. spicatum. Agropyron desertorum also had lower investment of nitrogen and biomass per unit area of photosynthetic tissues, more tillers and leaves per bunch, and shorter lived stems, all of which can contribute to greater tolerance of partial defoliation.Greater flexibility of resource allocation following defoliation was demonstrated by A. desertorum for both nitrogen and carbohydrates. Relatively more allocation to the shoot system and curtailed root growth in A. desertorum resulted in more rapid approach to the preclipping balance between the root and shoot systems, whereas root growth in A. spicatum continued unabated following defoliation. Nitrogen required for regrowth in both species was apparently supplied by uptake rather than reserve depletion. Carbohydrate pools in the shoot system of both species remained very low following severe defoliation and were approximately equivalent to carbon fixed in one day by photosynthesis of the whole canopy.Dedicated to Drs. Michael Evenari and Konrad Springer     

Nitrogen-15 partitioning within a three generation tiller sequence of the bunchgrass Schizachyrium scoparium: response to selective defoliation

Summary Nitrogen partitioning among three generations of tillers within the bunchgrass Schizachyrium scoparium var. frequens was investigated in a controlled environment as a potential mechanism of herbivory tolerance. Nitrogen-15 was transported from the labelled primary tiller generation to both shoots and roots of nondefoliated secondary and tertiary tiller generations within 24 h. Partial defoliation increased shoot nitrogen concentration of secondary and tertiary generation tillers by 110 and 120%, respectively, 24 h following defoliation. Shoot nitrogen concentration was preferentially increased by partial defoliation of tertiary generation tillers throughout the 120 h experimental period, but diminished to concentrations comparable to nondefoliated tillers within shoots of the secondary generation at 72 h. In contrast to nitrogen concentration, the total amount of nitrogen imported by secondary and tertiary generation tillers decreased 62 and 73%, respectively, 24 h following partial defoliation and did not attain values comparable to respective nondefoliated tillers. Consequently, preferential nitrogen concentration occurred in response to partial tiller defoliation without an increase in total nitrogen import based on the reduction in the total nitrogen requirement per tiller generation associated with defoliation. Estimates of both the total amount of nitrogen import and nitrogen concentration are necessary to accurately interpret the dynamics of intertiller nitrogen allocation.     

Root growth response to defoliation in two Agropyron bunchgrasses: field observations with an improved root periscope

Summary Root growth responses to defoliation were observed in the field with an improved root periscope technique, which is described. The grazing tolerant, Eurasian bunchgrass, Agropyron desertorum, was compared with the very similar but grazing sensitive, North American bunchgrass, A. spicatum. Root length growth of clipped A. desertorum was about 50% of that of intact plants, while root elongation of clipped A. spicatum continued relatively unabated during ninety days of regrowth following severe defoliation. The reduced root growth in A. desertorum was correlated with the allocation of relatively more resources to aboveground regrowth, thus aiding reestablishment of the root: shoot balance. This balance was apparent in similar root mortality patterns of clipped and control A. desertorum plants in the season following defoliation. In clipped A. spicatum, however, root mortality increased in the winter following the season in which the clipping was done and continued into the subsequent growing season. Reduction of root growth following defoliation appears to be an effective mechanism to aid reestablishment of the photosynthetic canopy and the root: shoot balance. As such it contributes to both herbivory tolerance and maintenance of competitive ability.     

Effects of bison grazing on Andropogon gerardii and Panicum virgatum in burned and unburned tallgrass paririe

Summary Responses to clipping and bison grazing in different environmental contexts were examined in two perennial grass species, Andropogon gerardii and Panicum virgatum, on the Konza Prairie in northeastern Kansas. Grazed tillers had lower relative growth rates (RGR) than clipped tillers following defoliation but this difference was transient and final biomass was not affected by mode of defoliation. Grazed tillers of both species had higher RGR throughout the season than ungrazed tillers, resulting in exact compensation for tissue lost to defoliation. However, A. gerardii tillers which had been grazed repeatedly the previous year (1988) had reduced relative growth rates, tiller biomass and tiller survival in 1989. This suggests that the short-term increase in aboveground relative growth rates after defoliation had a cost to future plant growth and tiller survival.In general, the two species had similar responses to defoliation but their responses were altered differentially by fire. The increase in RGR following defoliation of A. gerardii was relatively greater on unburned than burned prairie, and was influenced by topographic position. P. virgatum responses to defoliation were similar in burned and unburned prairie. Thus grazing, fire, and topographical position all interact to influence tiller growth dynamics and these two species respond differently to the fire and grazing interaction. In addition, fire may interact with grazing pattern to influence a plants' grazing history and thus its long-term performance.     

Demographic variability in tiller populations of two perennial pampa grasses

Abstract. The general objectives of this study were: (1) to investigate the importance of internal influences in regulating the tiller dynamics in natural populations of the warm-season perennial grasses Paspalum dilatatum and Sporobolus indicus, coexisting in Argentine flooded pampa, in as much as they act independently of the underlying external environment, and (2) to evaluate the extent to which interactions between internal and external factors affect the variation in tiller dynamics within such populations. Within-population variation in seasonal development of plants and tillers with different neighbour composition was studied for an annual growth cycle. Tiller survival and tillering were significantly influenced by tiller size. Tiller age influenced tiller fate, as suggested by the additive effects of age and size of tillers. These relationships varied with season and with species. Size and age of tillers showed additive effects with their neighbouring species on the tiller fate of P. dilatatum, but the effects of age and size of S. indicus changed according their neighbourhood. Tiller survival of S. indicus during the early growth season was more size-dependent when the cold-season species Poa lanigera, was the principal neighbour. Flowering and tillering probabilities were positively related through their common positive responses to tiller size. Tiller survival and recruitment between different seasons were strongly related. Independently of neighbour composition, tiller survival was generally inversely related to tiller recruitment in previous seasons. Therefore, significant density-dependent mortality of tillers was found for both species during the early summer when tiller density was expressed by basal area units.     

Does selective defoliation mediate competitive interactions in a semiarid savanna? A demographic evaluation

Abstract. Three patterns of target-neighbor plant defoliation were imposed on a late-seral, perennial, C₄-grass, Bouteloua curtipendula, in three long-term grazing regimes to determine the influence of selective defoliation on competitive interactions and species replacement in a semiarid savanna on the Edwards Plateau, Texas, USA. Short-term (3-yr) target plant defoliation did not significantly affect either tiller or plant responses in any of the three grazing regimes. Neighbor plant defoliation, either alone or in combination with target plants, produced a significant defoliation interaction with time for tiller number and basal area per plant, but not for tiller recruitment or mortality. The minimal effect of selective defoliation on the intensity of competitive interactions in this semiarid community indicates that selective grazing has a less definitive role in mediating herbivore-induced species replacement than it does in mesic grasslands and savannas. This interpretation is discussed within the context of long-term (45-yr) change in herbaceous vegetation associated with grazing in this community. Cumulative tiller recruitment in the intensively grazed regime was only 44% of that in the ungrazed regime because of greater plant mortality and fewer surviving plants that recruited tillers. Target plant mortality (50%) only occurred in the intensively grazed regime and the proportion of target plants that initiated tillers decreased by 70, 48 and 32% in the ungrazed, moderately and intensively grazed regimes, respectively, during the final two years of the investigation. The decrease in cumulative tiller recruitment in all grazing regimes was probably mediated by a drought-induced increase in median tiller age the second year of the study. However, tiller per tiller recruitment rate among plants that recruited at least one tiller remained relatively constant among grazing regimes and years. Intensive, long-term grazing has modified the population structure of this late-seral perennial grass to the extent that population responses to both herbivory and periodic drought have been altered in comparison with those of ungrazed and moderately grazed populations. Ecological consequences of a herbivore-induced transition in population structure may be to minimize the effect of selective herbivory on competitive interactions and to function as an avoidance mechanism to reduce the probability of localized population extinction in response to intensive long-term herbivory.     

Allocation of carbon to shoots,roots, soil and rhizosphere respiration by barrel medic (Medicago truncatula) before and after defoliation

The allocation of carbon to shoots, roots, soil and rhizosphere respiration in barrel medic (Medicago truncatulaGaertn.) before and after defoliation was determined by growing plants in pots in a labelled atmosphere in a growth cabinet. Plants were grown in a ¹⁴CO₂-labelled atmosphere for 30 days, defoliated and then grown in a ¹³CO₂-labelled atmosphere for 19 days. Allocation of ¹⁴C-labelled C to shoots, roots, soil and rhizosphere respiration was determined before defoliation and the allocation of ¹⁴C and ¹³C was determined for the period after defoliation. Before defoliation, 38.4% of assimilated C was allocated below ground, whereas after defoliation it was 19.9%. Over the entire length of the experiment, the proportion of net assimilated carbon allocated below ground was 30.3%. Of this, 46% was found in the roots, 22% in the soil and 32% was recovered as rhizosphere respiration. There was no net translocation of assimilate from roots to new shoot tissue after defoliation, indicating that all new shoot growth arose from above-ground stores and newly assimilated carbon. The rate of rhizosphere respiration decreased immediately after defoliation, but after 8 days, was at comparable levels to those before defoliation. It was not until 14 days after defoliation that the amount of respiration from newly assimilated C (¹³C) exceeded that of C assimilated before defoliation (¹⁴C). This revised version was published online in June 2006 with corrections to the Cover Date.     

Spatial arrangement of tiller replacement in Agropyron desertorum following grazing

Summary The spatial arrangement of tiller replacement was assessed on grazed and ungrazed tussocks of Agropyron desertorum (Fisch. ex Link) Schult. for three annual cycles. Frequency distributions of the number of replacement tillers per single progenitor were also determined. Tiller replacement was usually greater on the perimeter of tussocks than within the core, with or without grazing. Replacement was inversely related to grazing intensity, both on the perimeter and within the core of tussocks. Heights of replacement tillers on the perimeter or within the core seldom differed. Furthermore, grazing seldom affected the number of replacement tillers per progenitor. Greater tillering on the perimeter than within the core indicates that the tussocks were expanding. Apparently, grazing neither enhances tussock expansion and subsequent disintegration, nor does it necessarily lead to patches of tillers (multiple tillering per progenitor) within tussocks of A. desertorum.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The annual replacement of tillers of Agropyron desertorum (Fisch. ex Link) Schult., a grazing-tolerant, Eurasian tussock grass, was examined in the field following cattle grazing. Heavy grazing before internode (culm) elongation seldom affected tiller replacement. Heavy grazing during or after internode elongation, which elevates apical meristems, increased overwinter mortality of fall-produced tillers and reduced the number and heights of these replacement tillers. Unexpectedly, tussocks grazed twice within the spring growing season tended to have lower overwinter tiller mortality, greater tiller replacement, and larger replacement tillers than tussocks grazed only once in late spring. These responses of twice-grazed tussocks, however, were still less than those of ungrazed tussocks or tussocks grazed moderately in early spring. The presence of ungrazed tillers on partially grazed tussoks did not increase the replacement of associated grazed tillers relative to tillers on uniformly grazed plants. This result indicates that resource sharing among tillers, if present, is short-lived or ecologically unimportant in this species. Although A. desertorum is considered grazing-tolerant, tiller replacement on heavily grazed tussocks, particularly those grazed during or after internode elongation when apical meristems were removed, was usually inadequate for tussock maintenance. These observations at the tiller (ramet) level of organization in individual tussocks (genet) may explain the often noted reduction in stand (population) longevity with consistent heavy grazing.     

SEED DYNAMICS OF ABIES BALSAMEA AND ACER SACCHARUM IN A DECIDUOUS FOREST OF NORTHEASTERN NORTH AMERICA

We studied the seed dynamics of Abies balsamea and Acer saccharum in a sugar maple-yellow birch forest of Québec, Canada. Seed rain was censused every week from spring to autumn in 1988 and 1989. The soil seed bank was sampled four times during the 1988 growing season. Abies seed rain varied significantly between 1988 and 1989. A total of 92.5 ± 7.0 seeds nr^-2 (mean ± 1 SE) were produced in 1988; 1989 seed production was null. Seed viability in 1988 averaged 31.5%. Dispersal during the winter period accounted for ca. 22% of the annual seed crop. The spatial dispersion pattern of the seed rain was contagious for the autumn period, but was not significantly different from random for the winter period. There was only a weak relationship in the spatial distribution of the seed rain between the autumn and winter periods. Abies did not maintain a persistent seed bank on the site; viable seeds were present on the soil only after seed dispersal had started. Postdispersal seed mortality was relatively high, ca. 70%. Acer seed rain started soon after the flowering period, but consisted at that time only of aborted, underdeveloped samaras. Viable seeds were disseminated from September through early winter. Dispersal during the autumn accounted for ca. 98% of the annual seed crop. There were significant differences between seed rain abundance of 1988 and that of 1989 (118.1 ± 9.5 samaras m^-2 and 158.9 ± 16.3 samaras m^-2 in 1988 and in 1989, respectively). Seed viability also varied significantly between the 2 years, i.e., 19% in 1988 and 5% in 1989. For both 1988 and 1989, samaras were contagiously dispersed over the site. There was a significant positive relationship between the spatial distribution of the seed rain in 1988 and that in 1989. Seed rain abundance was high mainly in the proximity of mature Acer trees. As for Abies, Acer did not maintain a persistent seed bank in the soil; viable seeds were present only in the autumn seed bank sampling, after seed fall had started. Postdispersal seed mortality was relatively low, i.e., 20%. Abies and Acer are quite similar in their regenerative traits, and these contrast sharply with those of Betula alleghaniensis, a regular member of this forest community. Differences in regenerative traits may contribute to the coexistence of these tree species considering the gap regime of the system studied.     

Comparative demography of co-occurring introduced and native tussock grasses: persistence and potential expansion

Summary Demographic characteristics associated with the maintenance and growth of populations, such as seed dynamics, seedling emergence, survival, and tiller dynamics were examined for two tussock grasses, the native Agropyron spicatum and the introduced Agropyron desertorum in a 30-month field study. The introduced grass was aerially sown onto a native grassland site. Seed production of the introduced grass was greater than the native grass in both above- and below-average precipitation years. Seeds of A. spicatum were dispersed when they mature, while A. desertorum retained some seeds in inflorescences, and dispersed them slowly throughout the year. This seed retention allowed some seeds of the introduced grass to escape peak periods of seed predation during the summer and allowed seeds to be deposited constantly into the seed bank. Carryover of seeds in the seed bank beyond one year occurred in the introduced grass but not in the native species. For both species, seedling emergence occurred in both autumn or spring. Survival rates for A. desertorum were higher than A. spicatum when seedlings emerged between November and March. Survival rates of cohorts emerging before November favored A. spicatum whereas survival rates did not differ between species for cohorts emerging after March. Individuals of both species emerging after April were unable to survive the summer drought. Demographic factors associated with seeds of A. desertorum seemed to favor the maintenance and spread of this introduced grass into native stands formerly dominated by A. spicatum.     

Carbon import among vegetative tillers within two bunchgrasses: assessment with carbon-11 labelling

Summary Carbon allocation among bunchgrass tillers was examined with carbon-11 (¹¹CO₂) steady state labelling. Labelled carbon was continuously transported from parent tillers to anatomically attached daughter tillers at a time when morphological characteristics indicated that tiller maturation had occurred. Steady state levels of import into monitored daughter tillers increased within 30 min of either defoliation or shading. Import levels decreased within 30 min of the removal of shading, but remained accelerated throughout an 84 h observation period following defoliation. A second defoliation further increased carbon import into a monitored tiller above the previously accelerated level resulting from the initial defoliation. Carbon import by vegetative tillers in the two bunchgrass species examined may be most appropriately viewed as a series of potentially accelerated import levels above a low level of continuous import.     

Contribution of flexible allocation priorities to herbivory tolerance in C4 perennial grasses: an evaluation with 13C labeling

The ability of plants to rapidly replace photosynthetic tissues following defoliation represents a resistance strategy referred to as herbivory tolerance. Rapid reprioritization of carbon allocation to regrowing shoots at the expense of roots following defoliation is a widely documented tolerance mechanism. An experiment was conducted in a controlled environment to test the hypothesis that herbivory-sensitive perennial grasses display less flexibility in reprioritizing carbon allocation in response to defoliation than do grasses possessing greater herbivory tolerance. An equivalent proportion of shoot biomass (60% dry weight) was removed from two C₄ perennial grasses recognized as herbivory-sensitive, Andropogon gerardii and Schizachyrium scoparium, and two C₄ perennial grasses recognized as herbivory-tolerant, Aristida purpurea and Bouteloua rigidiseta. Both defoliated and undefoliated plants were exposed to ¹³CO₂ for 30 min, five plants per species were harvested at 6, 72 and 168 h following labeling, and biomass was analyzed by isotope ratio mass spectrometry. The tallgrass, A. geraiddii, exhibited inflexible allocation priorities while the shortgrass, B. rigidiseta, exhibited flexible allocation priorities in response to defoliation which corresponded with their initial designations as herbivory-sensitive and herbivory-tolerant species, respectively. A. gerardii had the greatest percentage and concentration of ¹³C within roots and lowest percentage of ¹³C within regrowth of the four species evaluated. In contrast, B. rigidiseta had a greater percentage of ¹³C within regrowth than did A. gerardii, the greatest percentage of ¹³C within new leaves of defoliated plants, and the lowest concentration of ¹³C within roots follwing defoliation. Although both midgrasses, S. scoparium and A. purpurea, demonstrated flexible allocation priorities in response to defoliation, they were counter to those stated in the initial hypothesis. The concentration of ¹³C within new leaves of S. scoparium increased in response to a single defoliation while the percentage and concentration of ¹³C within roots was reduced. A. purpurea was the only species in which the percentate of ¹³C within new leaves decreased while the percentage of ¹³C within roots increased following defoliation. The most plausible alternative hypothesis to explain the inconsistency between the demonstrated responsiveness of allocation priorities to defoliation and the recognized herbivory resistance of S. scoparium and A. purpurea is that the relative ability of these species to avoid herbivory may make an equal or greater contribution to their overall herbivory resistance than does herbivory tolerance. Selective herbivory may contribute to S. scoparium's designation as a herbivorysensitive species even though it possesses flexible allocation priorities in response to defoliation. Alternatively, the recognized herbivory resistance of A. purpurea may be a consequence of infrequent and/or lenient herbivory associated with the expression of avoidance mechanisms, rather than the expression of tolerance mechanisms. A greater understanding of the relative contribution of tolerance and avoidance strategies of herbivory resistance are required to accurately interpret how herbivory influences plant function, competitive interactions, and species abundance in grazed communities.     

Wolf predation and snow cover as mortality factors in the ungulate community of the Bialowieża National Park,Poland

Summary Wolf-ungulate interactions were studied in the pristine deciduous and mixed forests of the Bialowiea National Park in 1985–1989. The study period included two severe and two mild winters. The community of ungulates inhabiting Bialowiea National Park consisted of red deer Cervus elaphus, 55% of all ungulates; wild boar Sus scrofa, 42%; and roe deer Capreolus capreolus, moose Alces alces, and European bison Bison bonasus, about 1% each. The average size of red deer groups increased from 2.7 (SD 2.35) in spring and summer to 6.9 (SD 6.84) in autumn and winter. In winter the group size of red deer was positively correlated with the depth of snow cover and negatively correlated with the mean daily temperature. Average group size of wild boar did not change significantly between seasons; it was 6.8 (SD 5.16) in spring and summer and 5.7 (SD 4.67) in autumn and winter. Analysis of 144 wolf scats showed that wolves preyed selectively on red deer. In October–April, Cervidae (mostly red deer) constituted 91% of biomass consumed by wolves, while wild boar made up only 8%. In May–September deer formed 77% of prey biomass, and the share of wild boar increased to 22%. In all seasons of the year wolves selected juveniles from deer and boar populations: 61% of red deer and 94% of wild boar of determined age recovered from wolves' scats were young <1 year old. Analysis of 117 carcasses of ungulates found in Bialowiea National Park showed that predation was the predominant mortality factor for red deer (40 killed, 10 dead from causes other than predation) and roe deer (4 killed, none dead). Wild boar suffered most from severe winter conditions (8 killed, 56 dead). The percentage of ungulates that had died from undernutrition and starvation in the total mortality was proportional to the severity of winter.     

Defoliation and below-ground herbivory in the grass Muhlenbergiaquadridentata : Effects on plant performance and on the root-feeder Phyllophaga sp. (Coleoptera, Melolonthidae)

In this study we evaluated (1) the combined effects of simulated defoliation and below-ground herbivory (BGH) on the biomass and nitrogen content of tillers and roots of the bunchgrass Muhlenbergia quadridentata and (2) the effect of defoliation on the survival of third-instar root-feeder larvae of Phyllophaga sp. The experiment was performed in a pine forest area at an altitude of 3200 m above sea level. The grass and the root-feeder species were native and dominant in the understory and in the macroarthropod root-feeder communities, respectively. Plants were established in pots in the field and were subjected to the following treatments in a factorial design: simulated defoliation (three levels) and BGH (with or without root-feeder larvae) with ten replicates per treatment. Plants were defoliated three times at 2-month intervals. The interaction between defoliation and root herbivory was significant for all components of plant biomass. In every case, light defoliation with BGH decreased live above-ground, root and total plant biomass, and the number of live tillers by more than 50% with respect to the same defoliation level without root-feeders. Plants apparently did not compensate for the carbon drain by root-feeders when a high proportion of older leaves were not removed by defoliation. Plants under heavy defoliation were not affected by the presence of root-feeders and showed a greater live/dead above-ground biomass ratio than lightly defoliated and control plants. Defoliation and BGH did not change tiller and root N concentrations but root herbivores did decrease live-tiller N content in lightly defoliated plants. Root-feeders but not defoliation decreased the root/shoot ratio by 40% and the live/dead above-ground biomass ratio by 45% through increased tiller mortality. Survivorship and final biomass of Phyllophaga sp. larvae were not affected by defoliation treatments during the 6-month study period. Received: 17 May 1996 / Accepted: 1 November 1996     

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.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Drugs from emasculated hormones: The principle of syntopic antagonism

This lecture outlines the early stages in the discovery of adrenaline -receptor antagonists and of the histamine H₂-receptor antagonists. It ends with a brief personal view about future research.Published inLes Prix Nobel 1988, printed in Sweden by Norstedts Tryckeri, Stockholm, Sweden, 1989, republished here with the permission of the Nobel Foundation, the Copyright holder.     

Seasonal variations in the chemical composition of dissolved organic matter in organic forest floor layer leachates of old-growth Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) stands in northeastern Bavaria, Germany

Organic matter dissolved in thepercolation water of forest soils contributeslargely to element cycling and transport ofnatural and anthropogenic compounds. The wayand extent to which these processes areaffected depends on the amount and the chemicalcomposition of soluble organic matter. Becausethe amount of soluble organic matter variesseasonally with changes in the microbialactivity in soil, it seems reasonable to assumethat there may be also seasonal changes in thechemical composition of dissolved organicmatter. We examined dissolved organic matter inthe seepage waters of organic forest floorlayers over a 27-month period (1997–1999) intwo forest ecosystems, a 160-year-old Scotspine (Pinus sylvestris L.) stand and a90-year-old European beech (Fagussylvatica L.) forest. The forest floorleachates were analysed for bulk dissolvedorganic C, C in hydrophilic and hydrophobicdissolved organic matter fractions,lignin-derived phenols (CuO oxidation),hydrolysable neutral carbohydrates and uronicacids, hydrolysable amino sugars, and stablecarbon isotope composition. In addition, westudied the samples by use of liquid-state¹³C-nuclear magnetic resonance (NMR)spectroscopy.For both investigated forest sites we foundthat the dissolved organic carbonconcentrations in forest floor leachates werelargest during summer. They peaked after rainstorms following short dry periods (106–145 mgdissolved organic C l^–1). The proportionsof C in the hydrophilic fractions were largestin winter and spring whereas in summer andautumn more C was found in the hydrophobicfraction. According to liquid-state ¹³C-NMR spectroscopy, summer and autumn samples hadlarger abundances of aromatic and aliphaticstructures as well as larger proportions ofcarboxyl groups whereas the winter and springsamples were dominated by resonances indicatingcarbohydrates. Wet-chemical analyses confirmedthese results. Winter and spring samples wererich in neutral carbohydrates and amino sugars.The summer and autumn samples contained morelignin-derived phenols which were also strongeroxidised than those in the winter and springsamples. Seasonal changes of ¹³C valueswere found to reflect the changes in thechemical composition of dissolved organicmatter. Most negative values occurred whenisotopically light lignin-derived compoundswere abundant and less negative values whencarbohydrates predominated.The different vegetation, age of thestands, and underlying mineral soils resultedin different concentrations of dissolvedorganic carbon and in differences in thedistribution between hydrophobic andhydrophilic organic carbon. Despite of this,the results suggest that the trends in temporalvariations in the composition of dissolvedorganic matter in forest floor seepage waterwere remarkably similar for both sites.Dissolved organic matter in winter and springseems to be mainly controlled by leaching offresh disrupted biomass debris with a largecontribution of bacterial and fungal-derivedcarbohydrates and amino sugars. Dissolvedorganic matter leached from the forest floor insummer and autumn is controlled by thedecomposition processes in the forest floorresulting in the production of stronglyoxidised, water-soluble aromatic and aliphaticcompounds. The chemical composition ofdissolved organic matter in forest floorseepage water in winter and spring indicateslarger mobility, larger biodegradability, andless interaction with metals and organicpollutants than that released during summer andautumn. Thus, the impact of dissolved organicmatter on transport processes may varythroughout the year due to changes in itscomposition.     

Responses to simulated herbivory and water stress in two tropical C4 grasses

Summary The African grass Hyparrhenia rufa has established itself successfully in South American savannas (Llanos) and displaced dominant native grasses such as Trachypogon plumosus from the wetter and more fertile habitats. Several ecophysiological traits have been related to the higher competitive capacity of H. rufa. To further analyze the behavior of both species, their growth, biomass allocation, physiological and architectural responses to defoliation and water stress were compared under controlled conditions. Although total, aerial and underground biomass decreased under defoliation in both grasses, increases in clipped-leaf biomass and area compensated for defoliation in H. rufa but not in T. plumosus. This difference was due mainly to a higher proportion of assimilates being directed to leaf and tiller production and a higher leaf growth rate in the African grass as compared to T. plumosus, which showed incrased senescence under frequent defoliation. In both species, water stress ameliorated the effects of defoliation. The ability to compensate for defoliated biomass in H. rufa is possibly related to its long coevolution with large herbivores in its original African habitat and is apparently one of the causes of its success in Neotropical savannas.