Composition of soil Frankia assemblages across ecological drivers parallels that of nodule assemblages in Alnus incana ssp. tenuifolia in interior Alaska.

In root nodule symbioses (RNS) between nitrogen (N)-fixing bacteria and plants, bacterial symbionts cycle between nodule-inhabiting and soil-inhabiting niches that exert differential selection pressures on bacterial traits. Little is known about how the resulting evolutionary tension between host plants and symbiotic bacteria structures naturally occurring bacterial assemblages in soils. We used DNA cloning to examine soil-dwelling assemblages of the actinorhizal symbiont Frankia in sites with long-term stable assemblages in Alnus incana ssp. tenuifolia nodules. We compared: (1) phylogenetic diversity of Frankia in soil versus nodules, (2) change in Frankia assemblages in soil versus nodules in response to environmental variation: both across succession, and in response to long-term fertilization with N and phosphorus, and (3) soil assemblages in the presence and absence of host plants. Phylogenetic diversity was much greater in soil-dwelling than nodule-dwelling assemblages and fell into two large clades not previously observed. The presence of host plants was associated with enhanced representation of genotypes specific to A. tenuifolia, and decreased representation of genotypes specific to a second Alnus species. The relative proportion of symbiotic sequence groups across a primary chronosequence was similar in both soil and nodule assemblages. Contrary to expectations, both N and P enhanced symbiotic genotypes relative to non-symbiotic ones. Our results provide a rare set of field observations against which predictions from theoretical and experimental work in the evolutionary ecology of RNS can be compared.

Experimental evidence that ptarmigan regulate willow bud production to their own advantage.

In some ecosystems, vertebrate herbivores increase the nutritional quality and biomass of their food source through repeated grazing, thereby manipulating their environment to support higher densities of animals. We tested whether ptarmigan (Lagopus lagopus and L. muta) are capable of regulating the nutritional quality, abundance, and availability of feltleaf willow (Salix alaxensis) buds using a simulated browsing experiment and a feeding preference study with wild birds. Simulated ptarmigan browsing resulted in smaller buds, but greater numbers of buds per shoot. Furthermore, browsing altered the morphology of willow branches such that buds were at higher densities and closer to snow level compared to unbrowsed controls. Browsing increased the number of willows with accessible buds (buds within 50 cm of snow level) from 55 to 89%, and increased total accessible bud biomass from 113 ± 30 to 129 ± 50 mg/ramet. Browsing did not affect bud nitrogen or carbon concentration and slightly reduced protein precipitation capacity (tannins) in buds the following winter, indicating that ptarmigan browsing does not induce a defensive response in this species. When branches of broomed (previously browsed) and unbroomed willows were placed in the snow at equal heights, ptarmigan showed no preference for either type; however, they obtained more buds from broomed willows. Increased accessibility and density of willow buds caused by browsing has the potential to increase habitat carrying capacity, thereby supporting higher densities of ptarmigan.

Phylogeny and assemblage composition of Frankia in Alnus tenuifolia nodules across a primary successional sere in interior Alaska.

In nitrogen (N) fixing symbioses, host-symbiont specificity, genetic variation in bacterial symbionts and environmental variation represent fundamental constraints on the ecology, evolution and practical uses of these interactions, but detailed information is lacking for many naturally occurring N-fixers. This study examined phylogenetic host specificity of Frankia in field-collected nodules of two Alnus species (A. tenuifolia and A. viridis) in interior Alaska and, for A. tenuifolia, distribution, diversity, spatial autocorrelation and correlation with specific soil factors of Frankia genotypes in nodules collected from replicated habitats representing endpoints of a primary sere. Frankia genotypes most commonly associated with each host belonged to different clades within the Alnus-infective Frankia clade, and for A. tenuifolia, were divergent from previously described Frankia. A. tenuifolia nodules from early and late succession habitats harboured distinct Frankia assemblages. In early succession, a single genotype inhabited 71% of nodules with no discernable autocorrelation at any scale, while late succession Frankia were more diverse, differed widely among plants within a site and were significantly autocorrelated within and among plants. Early succession Frankia genotype occurrence was strongly correlated with carbon/nitrogen ratio in the mineral soil fraction, while in late succession, the most common genotypes were correlated with different soil variables. Our results suggest that phylogenetic specificity is a significant factor in the A. tenuifolia-Frankia interaction and that significant habitat-based differentiation may exist among A. tenuifolia-infective genotypes. This is consistent with our hypothesis that A. tenuifolia selects specific Frankia genotypes from early succession soils and that this choice is attenuated in late succession.

The role of lesser snow geese as nitrogen processors in a sub-arctic salt marsh.

Ammonia volatilization losses from faeces of Lesser Snow Geese were measured during the summer of 1987 on the salt-marsh flats at La Pérouse Bay. Amounts of ammonia volatilized increased with increasing ambient temperature, and ranged from 1.0 to 15.1 mg N per 100 mg of nitrogen present as soluble ammonium ions at the start of the 8-h experiment. Using estimates of faecal deposition reported previously, the annual loss via volatilization was estimated at 0.08 g N m-2, or 7.9% of the nitrogen present in goose faeces. Percent change in soluble ammonium ions in fresh faeces after 8 h ranged from -51.1% to +41.1%, indicating that net mineralization of organic nitrogen occurred in some of the faeces. Microbial respiration of fresh goose faeces increased exponentially with temperature. However, variable rates of net mineralization per unit rate of respiration indicated that the substrate quality affected microbial immobilization and thus net nitrogen mineralization. In feeding experiments, captive goslings grazed different types of vegetation, each with distinctive nutritional qualities. Forage quality had significant effects on goose feeding behavior and subsequent rates of nitrogen mineralization in fresh faeces. Net nitrogen mineralization rates in faeces from geese which grazed the three vegetation types ranged from 1.31 to 4.97 mg NH 4+ -N gDW-1 24 h-1. Because plant growth in this salt marsh is nitrogen-limited, where swards are grazed, mineralization of organic faecal nitrogen represents an essential link in the maintenance of the flow of nitrogen into the sediments and the sustained growth of vegetation at a time when most required by the geese.

The interaction of defoliation and nutrient uptake in Sporobolus kentrophyllus, a short-grass species from the serengeti plains.

Sporobolus kentrophyllus, a grazing-tolerant C4 grass from the southeastern Serengeti Plains, was grown in solution culture to examine the effects of clipping on the uptake, preference and subsequent transport of varying nitrogen forms. Clipping reduced offtake mass, crown mass ane root mass, resulting in a 58% decline in plant mass. Proportional biomass allocation to roots decreased with clipping, while tillering rates increased. Clipping also increased the nitrogen concentrations of all tissues, and plant nitrogen uptake (nitrogen accumulated throughout the experiment per gram root). The 15N concentrations (% atom excess) of all tissues were higher in clipped compared with unclipped plants, and the average 15N uptake rate of clipped plants was twice that of unclipped plants. The relative 15N allocation to aboveground mass, a measure of canopy sink strength, was higher in clipped plants. Plants fed 15N-ammonium or 15N-nitrate during the 15N pulse experiment had greater 15N tissue concentrations compared with urea-fed plants, and 15N uptake rates were higher in ammonium-fed and nitrate-fed plants, compared with urea-fed plants. The relative magnitudes of these differences were higher when plants were clipped. Clipped plants had higher uptake rates for potassium, phosphorus and sodium, while differences between clipping treatments for calcium, iron, and magnesium were indistinguishable. Rapid uptake rates for species on the southeastern Serengeti plains, particularly during grazing periods, have important implications for nutrient cycling in this system.

Ammonia volatilization and the effects of large grazing mammals on nutrient loss from East African grasslands.

Ammonia volatilization losses measured from soils at seven sites in the Serengeti National Park, Tanzania during the 1986 growing season ranged from 2.78±0.49% to 25.03±1.34% of nitrogen applied. Although peak ammonia losses ranged from 0.071±0.018 to 0.404±0.040 g N m-2 h-1, rates dropped to zero within four days, and calculations reveal that volatilization losses represent minor fluxes in the context of the system's nitrogen cycling. Volatilization losses were inversely correlated with grazing intensity experienced by a site, and it appears that large ungulates themselves contribute to nutrient conservation throught indirect interactive effects on system processes.

Urea as a promotive coupler of plant-herbivore interactions.

Growth responses of Kyllinga nervosa Steud., a sedge from the Serengeti short-grass plains, were examined in a factorial experiment which included clipped and unclipped plants, and nitrogen supplied as either urea or ammonium nitrate. Results were expressed in relation to three transfer processes: flow to grazers, flow to producers and flow to reproduction. Clipping increased biomass and nitrogen flow to grazers by significantly increasing nitrogen uptake, aboveground nitrogen flow, and the weights of and proportional allocation to green leaf production. This was at the expense of flow to vegetative and sexual reproduction, since the weights and proportional investments in roots, crowns and reproductive structures were reduced. Urea nutrition increased flow to grazers and plant reproduction through increases in green leaf weight, flower weight, allocation to green leaves, flowers and stems, and aboveground: belowground biomass ratios. Stimulation of aboveground productivity by urea was a consequence of increased tillering rates.Interactive responses of clipping and nitrogen source regulated plant growth, thus controlling flow to each transfer process. Combined effects of clipping and urea resulted in compensatory production of both green leaves and flowers, and maximized biomass and nitrogen flow to grazers. Both urea and clipping tightened herbivore-producer recycling by significantly reducing litter nitrogen and carbon masses. In contrast, when plants were unclipped and grown on NH4NO3, biomass allocation and weights of roots and crowns were increased at the expense of aboveground tissues, thus increasing flow to primary producers. Plant growth responses to experimental treatment combinations simulating nutritional status of grazed and ungrazed field plants indicate that urea represents a potential importance beyond it nitrogen contribution by introducing a positive feedback to herbivores.

The effects of clipping, nitrogen source and nitrogen concentration on the growth responses and nitrogen uptake of an east african sedge.

Kyllinga nervosa Steud., a sedge from the Serengeti short-grass plains, was subjected to a balanced factorial experiment which included unclipped plants and plants clipped weekly to a 5 cm height, nitrogen supplied as either nitrate or ammonium and three nitrogen concentrations. Tillering rates, green leaf nitrogen, and both green leaf weight and biomass investment in green leaf production increased with nitrogen concentration. Low nitrogen conserved investment in crown production and resulted in adjustments for nitrogen acquisition by increasing biomass allocation to root production. Nitrate nutrition stimulated green leaf weight, tillering rate, nitrogen redistribution and both crown and root nitrogen. Ammonium nutrition increased nitrogen uptake, total plant nitrogen accumulation, reproduction, litter weight and nitrogen loss to decomposers. Clipping increased investment in green leaf production at the expense of stem, root, crown and flower production. Compensatory green leaf production in response to clipping occurred only when plants were grown in ammonium. Clipping stimulated uptake rates of both ammonium and nitrate, and therefore total plant nitrogen accumulation. Results suggest a balanced utilization of both nitrate and ammonium may be necessary for optimal growth in this species.