Resource allocation patterns of two California-Sonoran desert ephemerals

Summary The patterns of allocation of structural and nonstructural carbon were followed in the co-occurring desert ephemerals Plantago insularis and Camissonia boothii. Patterns of biomass distribution were determined from material harvested at biweekly intervals as were levels of nonstructural sugar and starch. Seasonal patterns of growth and reproduction differed markedly with Plantago allocating significantly more structural and nonstructural carbon to reproduction early in the season. Plantago completed its life cycle in less than 60 days but Camissonia continued both vegetative and reproductive growth to over 100 days. The longer growing season of Camissonia was possible because more energy was allocated to vegetative tissues and storage presumably as investment toward longer life and higher levels of reproduction.     

Circumpolar arctic tundra biomass and productivity dynamics in response to projected climate change and herbivory

Satellite remote sensing data have indicated a general ‘greening’ trend in the arctic tundra biome. However, the observed changes based on remote sensing are the result of multiple environmental drivers, and the effects of individual controls such as warming, herbivory, and other disturbances on changes in vegetation biomass, community structure, and ecosystem function remain unclear. We apply ArcVeg, an arctic tundra vegetation dynamics model, to estimate potential changes in vegetation biomass and net primary production (NPP) at the plant community and functional type levels. ArcVeg is driven by soil nitrogen output from the Terrestrial Ecosystem Model, existing densities of Rangifer populations, and projected summer temperature changes by the NCAR CCSM4.0 general circulation model across the Arctic. We quantified the changes in aboveground biomass and NPP resulting from (i) observed herbivory only; (ii) projected climate change only; and (iii) coupled effects of projected climate change and herbivory. We evaluated model outputs of the absolute and relative differences in biomass and NPP by country, bioclimate subzone, and floristic province. Estimated potential biomass increases resulting from temperature increase only are approximately 5% greater than the biomass modeled due to coupled warming and herbivory. Such potential increases are greater in areas currently occupied by large or dense Rangifer herds such as the Nenets‐occupied regions in Russia (27% greater vegetation increase without herbivores). In addition, herbivory modulates shifts in plant community structure caused by warming. Plant functional types such as shrubs and mosses were affected to a greater degree than other functional types by either warming or herbivory or coupled effects of the two.     

Performance and allocation patterns of the perennial herb,Plantago lanceolata,in response to simulated herbivory and elevated CO2 environments

Summary We tested the prediction that plants grown in elevated CO₂ environments are better able to compensate for biomass lost to herbivory than plants grown in ambient CO₂ environments. The herbaceous perennial Plantago lanceolata (Plantaginaceae) was grown in either near ambient (380 ppm) or enriched (700 ppm) CO₂ atmospheres, and then after 4 weeks, plants experienced either 1) no defoliation; 2) every fourth leaf removed by cutting; or 3) every other leaf removed by cutting. Plants were harvested at week 13 (9 weeks after simulated herbivory treatments). Vegetative and reproductive weights were compared, and seeds were counted, weighed, and germinated to assess viability.Plants grown in enriched CO₂ environments had significantly greater shoot weights, leaf areas, and root weights, yet had significantly lower reproductive weights (i.e. stalks + spikes + seeds) and produced fewer seeds, than plants grown in ambient CO₂ environments. Relative biomass allocation patterns further illustrated differences in plants grown in ambient CO₂ environments. Relative biomass allocation patterns further illustrated differences in plant responses to enriched CO₂ atmospheres: enriched CO₂-grown plants only allocated 10% of their carbon resources to reproduction whereas ambient CO₂-grown plants allocated over 20%. Effects of simulated herbivory on plant performance were much less dramatic than those induced by enriched CO₂ atmospheres. Leaf area removal did not reduce shoot weights or reproductive weights of plants in either CO₂ treatment relative to control plants. However, plants from both CO₂ treatments experienced reductions in root weights with leaf area removal, indicating that plants compensated for lost above-ground tissues, and maintained comparable levels of reproductive output and seed viability, at the expense of root growth.     

Effects of simulated increased grazing on carbon allocation patterns in a high arctic mire

Herbivory is an important part of most ecosystems, and grazing alone can have a considerable impact on the ecosystems carbon balance with both direct and indirect effects. Removal of above-ground biomass by consumption of herbivores will change the below-ground carbon stock; the reduction of litter that goes into the ground will influence the total ecosystem carbon content. Little is however known about how plant-herbivory interactions effect the carbon balance, in particular methane emissions, of high arctic mires. We hypothesized that increased grazing pressure will change carbon allocation patterns resulting in decreased net ecosystem uptake of carbon and subsequently in lower methane emissions. An in-situ field experiment was conducted over 3 years in a high arctic mire at Zackenberg in NE Greenland. The experiment consisted of three treatments, with five replicates of each (1) control, (2) vascular plants were removed (NV), (3) clipped twice each growing season in order to simulate increased muskox grazing. Immediately after the initiation of the experiment net ecosystem uptake of CO₂ decreased in clipped plots (mean total decrease for the three following years was 35 %). One year into the experiment a significantly lower CH₄ emission was observed in these plots, the total mean reduction for the following 2 years was 26 %. Three years into the experiment significantly lower substrate (acetic acid) availability for CH₄ production was observed (27 % reduction). NV plots had a mean decrease in CO₂ uptake of 113 %, a 62 % decrease in ecosystem respiration and an 84 % decrease in CH₄ emission (mean of all 3 years). Our study shows that increased grazing pressure in a high arctic mire can lead to significant changes in the carbon balance, with lower CO₂ uptake leading to lower production of substrate for CH₄ formation and in lower CH₄ emission.     

Resource utilization and sex allocation in response to host size in two ectoparasitoid wasps on subcortical beetles

Patterns of host resource utilization and sex ratio manipulation in relation to host size were investigated for two solitary ectoparasitoid wasps,Atanycolus initiator andSpathius brevicaudis (Hymenoptera Braconidae). Both species parasitize subcortical beetles on the trunks of Japanese pine trees.A. initiator is on average 8 times larger in body weight and has an ovipositor that is 3.7 times longer than that ofS. brevicaudis. In both parasitoids, the size of emerging wasps was positively correlated with host size, but the host/wasp size regressions were linear for all three major host species inA. initiator, whereas inS. brevicaudis the regression was logarithmic for a relatively large host species. The sex ratios (proportion of males) of both parasitoids emerging from different host species decreased with increasing host size, but the overall sex ratio on each host species was male-biased inA. initiator, while female-biased inS. brevicaudis. How the proportion of host consumed changed in response to host size, differed between the two parasitoids for the same host species. In the field survey, the size and sex ratio of the emerging two parasitoids from a dead tree were closely related to host size. However, the spatial distribution of the two parasitoids depended on the bark thickness of the trunk. The data suggest that differences in the relative evaluation of host size and in ovipositor length may enable the coexistence of the two parasitoid wasps.     

Physiognomic changes in response to herbivory increase carbon allocation to roots in a temperate grassland of central Argentina

Plant Ecology - In most temperate grasslands, cattle grazing can promote physiognomic changes on plant communities, as well as changes in species growth patterns. Through these changes in...     

Aboveground resource allocation in response to root herbivory as affected by the arbuscular mycorrhizal symbiosis

Aims

Arbuscular mycorrhizal (AM) fungi associate with the majority of terrestrial plants, influencing their growth, nutrient uptake and defence chemistry. Consequently, AM fungi can significantly impact plant-herbivore interactions, yet surprisingly few studies have investigated how AM fungi affect plant responses to root herbivores. This study aimed to investigate how AM fungi affect plant tolerance mechanisms to belowground herbivory.

Methods

We examined how AM fungi affect plant (Saccharum spp. hybrid) growth, nutrient dynamics and secondary chemistry (phenolics) in response to attack from a root-feeding insect (Dermolepida albohirtum).

Results

Root herbivory reduced root mass by almost 27%. In response, plants augmented investment in aboveground biomass by 25%, as well as increasing carbon concentrations. The AM fungi increased aboveground biomass, phosphorus and carbon. Meanwhile, root herbivory increased foliar phenolics by 31% in mycorrhizal plants, and increased arbuscular colonisation of roots by 75% overall. AM fungi also decreased herbivore performance, potentially via increasing root silicon concentrations.

Conclusions

Our results suggest that AM fungi may be able to augment plant tolerance to root herbivory via resource allocation aboveground and, at the same time, enhance plant root resistance by increasing root silicon. The ability of AM fungi to facilitate resource allocation aboveground in this way may be a more widespread strategy for plants to cope with belowground herbivory.

     

Resource allocation of chilean herbs in response to climatic and microclimatic factors

Summary The standard metabolic rate (SMR) of birds correlates broadly with climate of origin. SMR tends to be higher in birds from cold climates and lower in tropical forms than would be expected from the bird's mass. SMR changes, on the average, 1% per degree change in latitude. The influence of climate on SMR is, however, subject to modification by other aspects of the bird's life history. For example, in tropical species adaptive modifications in SMR correlate with thermal microhabitat. Tropical birds which forage in the sun have SMR's averaging 25% lower than expected, while SMR of species which forage in the shade is normal. Species of penguins which undergo prolonged fasts during the breeding season do not show elevated SMR's typical of high latitude birds.     

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     

Productivity–diversity patterns in arctic tundra vegetation

Productivity has long been argued to be a major driver of species richness patterns. In the present study we test alternative productivity–diversity hypotheses using vegetation data from the vast Eurasian tundra. The productivity–species pool hypothesis predicts positive relationships at both fine and coarse grain sizes, whereas the productivity–interaction hypothesis predicts unimodal patterns at fine grain size, and monotonic positive patterns at coarse grain size. We furthermore expect to find flatter positive (productivity–species pool hypothesis) or more strongly negative (productivity–interaction hypothesis) relationships for lichens and bryophytes than for vascular plants, because as a group, lichens and bryophytes are better adapted to extreme arctic conditions and more vulnerable to competition for light than the taller‐growing vascular plants. The normalised difference vegetation index (NDVI) was used as a proxy of productivity. The generally unimodal productivity–diversity patterns were most consistent with the productivity–interaction hypothesis. There was a general trend of decreasing species richness from moderately to maximally productive tundra, in agreement with an increasing importance of competitive interactions. High richness of vascular plants and lichens occurred in moderately low productive tundra areas, whereas that of bryophytes occurred in the least productive tundra habitats covered by this study. The fine and coarse grain richness trends were surprisingly uniform and no variation in beta diversity along the productivity gradient was seen for vascular plants or bryophytes. However, lichen beta diversity varied along the productivity gradient, probably reflecting their sensitivity to habitat conditions and biotic interactions. Overall, the results show evidence that productivity–diversity gradients exist in tundra and that these appear to be largely driven by competitive interactions. Our results also imply that climate warming‐driven increases in productivity will strongly affect arctic plant diversity patterns.     

Aboveground biomass allocation,leaf growth,and photosynthesis patterns in tundra plant forms in arctic Alaska

Summary Tundra plant growth forms can generally be characterized as consisting predominantly of low-growing perennial grasses and sedges, perennial herbaceous forbs, dwarf deciduous shrubs, and dwarf evergreen shrubs. Gross aboveground carbon allocation, leaf growth, and photosynthesis pattern studies were initiated to develop a quantitative understanding of the functional importance of these particular tundra growth forms. Photosynthetic capacities of 13 species were determined under standardized exposure conditions using a¹⁴CO₂ field system and ranged between 5 and 47 mg CO₂·g dry wt^-1·h^-1. These results, in conjunction with detailed leaf growth determinations, support the generalization that species with an evergreen growth form have lower photosynthetic capacities than species with a perennial graminoid, forb, or deciduous shrub growth form. However, these low photosynthetic capacities in evergreen shrubs are associated with relatively extended leaf longevities. Conversely, deciduous shrub forms exhibited high photosynthetic capacities, but were offset by relatively short leaf longevity periods. The perennial grasses, sedges, and forbs showed patterns intermediate to these. As a result, it appears that among tundra species of different growth form, photosynthetic capacity is inversely related to leaf longevity.     

Geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata

We investigated geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata, to test the hypothesis that escape from herbivory in invasive species permits enhanced growth and lower production of defensive chemicals. We quantified herbivore damage, concentrations of sinigrin, and growth and reproduction inside and outside herbivore exclusion treatments, in field populations in the native and invasive ranges. As predicted, unmanipulated plants in the native range (Hungary, Europe) experienced greater herbivore damage than plants in the introduced range (Massachusetts and Connecticut, USA), providing evidence for enemy release, particularly in the first year of growth. Nevertheless, European populations had consistently larger individuals than US populations (rosettes were, for example, eightfold larger) and also had greater reproductive output, but US plants produced larger seeds at a given plant height. Moreover, flowering plants showed significant differences in concentrations of sinigrin in the invasive versus native range, although the direction of the difference was variable, suggesting the influence of environmental effects. Overall, we observed less herbivory, but not increased growth or decreased defense in the invasive range. Geographical differences in performance and leaf chemistry appear to be due to variation in the environment, which could have masked evolved differences in allocation.     

Resource allocation to reproduction in animals

The standard Dynamic Energy Budget (DEB) model assumes that a fraction κ of mobilised reserve is allocated to somatic maintenance plus growth, while the rest is allocated to maturity maintenance plus maturation (in embryos and juveniles) or reproduction (in adults). All DEB parameters have been estimated for 276 animal species from most large phyla and all chordate classes. The goodness of fit is generally excellent. We compared the estimated values of κ with those that would maximise reproduction in fully grown adults with abundant food. Only 13% of these species show a reproduction rate close to the maximum possible (assuming that κ can be controlled), another 4% have κ lower than the optimal value, and 83% have κ higher than the optimal value. Strong empirical support hence exists for the conclusion that reproduction is generally not maximised. We also compared the parameters of the wild chicken with those of races selected for meat and egg production and found that the latter indeed maximise reproduction in terms of κ, while surface‐specific assimilation was not affected by selection. We suggest that small values of κ relate to the down‐regulation of maximum body size, and large values to the down‐regulation of reproduction. We briefly discuss the ecological context for these findings.     

Composition and distributional patterns in arctic rotifers

Based on collections of rotifers from 212 localities in arctic North America, the patterns of distribution and composition are evaluated. An attempt is made to discern the dominant components of the rotifer community in arctic habitats. One hundred and sixty five species of rotifers are reported, three of which represent new records for North America. With increasing latitude and decreasing summer temperatures, a decline in species richness and change in species composition is observed. Some rotifers that previously were not adequately described are redescribed and illustrated. The significance of the dispersal capacity of rotifers in the arctic is discussed.     

In situ patterns of intracellular photosynthate allocation by sea ice algae in the Canadian high arctic

Summary A novel sampling/incubating device was used to determine the in situ patterns of intracellular photosynthate allocation by algae living in the bottom of annual sea ice in the Canadian arctic. During the seasonal decline of the bloom in late May and June, the average allocation pattern after 24 h incubation in the bottom 1 cm of ice (where the bulk of the algae are found) was 30.5% to low molecular weight materials, 10.6% to lipid, 48.8% to polysaccharide and 8.8% to protein. Allocation patterns were vertically stratified and light-dependent within the bottom ice community, with higher allocation to lipid in the upper, better-illuminated, strata. Chlorophyll-specific photosynthesis rates were extremely low (<0.031 gC·g Chl a ^–1·h^–1). Short incubations (ca. 1h) gave similar results to the 24 h incubations. The in situ allocation patterns were atypical of those normally expected for light-limited microalgae, but were consistent with a physiological response to inorganic nutrient limitation in the late stages of the bloom.     

Age‐related parasite load and longevity patterns in the sedge warbler Acrocephalus schoenobaenus

We report the results from a nine‐year study on parasite infection in males of a small migrant passerine, the sedge warbler Acrocephalus schoenobaenus. Every year, for each male caught during territory establishment we estimated infection intensity of two lineages of Haemoproteus belopolskyi (SW1 and SW3), using molecular methods. We found a significant relationship between infection intensity and males’ longevity in both studied lineages. There was severe mortality of second‐year sedge warblers after their first breeding season, with lower parasite load in survivors. The lower infection intensity in older age classes was related to between‐individual change in both lineages, but was also a result of differences in infection intensity during the lifetime of individual males in the SW3. The relation between parasite load and longevity suggests that parasite load may be an age‐dependent factor influencing individual survival.     

Seasonal temperature patterns in an arctic and two subarctic Alaskan (USA) headwater streams

We investigated the thermal ecology of three Alaskan streams. Monument Creek (MC) and Little Poker Creek (LPC) are subarctic streams in interior Alaska; LPC is in a permafrost-dominated valley. Imnavait Creek (IC) is an arctic tundra beaded stream in the northern foothills of the Brooks Range. Water temperatures were recorded with automated dataloggers hourly (LPC) or bi-hourly (MC and IC). Records for MC extend through almost three entire years, while data from IC (three years) and LPC (one year) represent the majority of the ice-free season. We also collected winter water/ice temperatures from IC (1989–1990). Mean annual water temperatures were 1.1 °C (LPC), 2.3 °C (MC), and 2.9 °C (IC), while maxima were 5.8 °C (LPC), 13.0 °C (MC), and 21.4 °C (IC). Water temperature rose in the spring about twice as fast (both mean and maximum daily increase) in MC as in LPC, and again about twice as fast in IC as in MC. A similar pattern was observed during the autumnal decline in water temperature. Maximum daily amplitude followed a similar pattern, with MC (6.6 °C) intermediate between LPC (4.1°) and IC (11.6°). LPC accumulated approximately 400 degree-days above 0 °C, MC approximately 950 degree-days, and IC approximately 1000 degree-days. Although it is about 450 km north of the other streams, the tundra stream (IC) accumulated more degree-days, had higher maximum and mean temperatures, greater daily temperature amplitude, and steeper slopes of vernal temperature rise and autumnal temperature decline than the subarctic streams (LPC and MC). The absence of a canopy of riparian plants, channel morphology, and continuous sunlight during the arctic mid-summer accounted for these higher temperatures. Beaded tundra streams provide a highly seasonal (< 120 d ice-free) and spatially and temporally complex thermal environment.