Soil carbon dynamics: the effects of nitrogen input, intake demand and off-take by animals.

Elucidation of the drivers of soil carbon (C) change is required to enable decisions to be made on how to achieve soil C sequestration. Interactions between different components in the ecosystem in combination with feedback mechanisms mean that identifying drivers through conventional experimental approaches or by retro-fitting models to data are unlikely to result in the insights needed for the future. This paper explains soil C dynamics by using a process-based model. Drivers considered in the model include nitrogen (N) fertiliser inputs, intake demand, and off-take of animal products. The effect of the grazing animal in uncoupling the C and N cycles is explained, plus the implications of the farming system ('drystock' versus milk). The model enables depiction of the dynamic equilibrium achieved with time when a proposed change in the drivers is sustained. The results show that soil C loss under lactating cows is a result of N, rather than C, being removed in milk. Counter-intuitively, at the same intake demand, N loss under 'milk' is less than under 'dry-stock', as is C loss in animal respiration. Possibilities for changing the longevity of C in the soil are discussed, and the compromise between food production, N loss and C sequestration is considered.

Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2.

Atmospheric CO2 enrichment may stimulate plant growth directly through (1) enhanced photosynthesis or indirectly, through (2) reduced plant water consumption and hence slower soil moisture depletion, or the combination of both. Herein we describe gas exchange, plant biomass and species responses of five native or semi-native temperate and Mediterranean grasslands and three semi-arid systems to CO2 enrichment, with an emphasis on water relations. Increasing CO2 led to decreased leaf conductance for water vapor, improved plant water status, altered seasonal evapotranspiration dynamics, and in most cases, periodic increases in soil water content. The extent, timing and duration of these responses varied among ecosystems, species and years. Across the grasslands of the Kansas tallgrass prairie, Colorado shortgrass steppe and Swiss calcareous grassland, increases in aboveground biomass from CO2 enrichment were relatively greater in dry years. In contrast, CO2-induced aboveground biomass increases in the Texas C3/C4 grassland and the New Zealand pasture seemed little or only marginally influenced by yearly variation in soil water, while plant growth in the Mojave Desert was stimulated by CO2 in a relatively wet year. Mediterranean grasslands sometimes failed to respond to CO2-related increased late-season water, whereas semiarid Negev grassland assemblages profited. Vegetative and reproductive responses to CO2 were highly varied among species and ecosystems, and did not generally follow any predictable pattern in regard to functional groups. Results suggest that the indirect effects of CO2 on plant and soil water relations may contribute substantially to experimentally induced CO2-effects, and also reflect local humidity conditions. For landscape scale predictions, this analysis calls for a clear distinction between biomass responses due to direct CO2 effects on photosynthesis and those indirect CO2 effects via soil moisture as documented here.

Relationships among shoot sinks for resources exported from nodal roots regulate branch development of distal non-rooted portions of Trifolium repens L.

Two manipulative experiments tested hypotheses pertaining to the correlative control exerted by nodal roots on branch development of the distal non-rooted portion of Trifolium repens growing clonally under near-optimal conditions. The two experiments, differing in their pattern of excision to manipulate the number of branches formed at the first 9-10 phytomers distal to the youngest nodal root, each found that after 20 phytomers of growth the total number of lateral branches formed on the primary stolon remained between five and seven regardless of where the branches formed along the stolon. Additional treatments established that nodal roots influenced branch development via relationships among shoot sinks for the root-supplied resources rather than through variation in the supply of such resources induced by fluctuations in photosynthate supply to roots from branches. Regression analysis of data pooled from treatments of both experiments confirmed that shoot-sink relationships for root- supplied resources controlled the branching processes on the non-rooted portion of plants. A disbudding treatment, which removed all the apical and axillary buds present on basal branches, but left other branch tissues intact, increased branch development of the apical region in the same way as did complete excision of the basal lateral branches. The apical buds and the elongation processes occurring immediately proximal to the buds were thus identified as strong sinks for the root-supplied resources. Such results suggest that branch development on the non-rooted shoot portion distal to the youngest nodal root is regulated by competition among sinks for root-derived resources, of limited availability, necessary for the processes of elongation of axillary buds and the primary stolon apical bud.

A developmentally based categorization of branching in Trifolium repens L.: influence of nodal roots.

This study describes the successive stages of development of branches from axillary buds in fully rooted plants of Trifolium repens grown in near optimal conditions, and the way in which this developmental pathway differs when nodal root formation is prevented as plants grow out from a rooted base. Cuttings of a single genotype were established in a glasshouse with nodal root systems on the two basal phytomers and grown on so that nodal rooting was either permitted (+R) or prevented (-R). In +R plants, axillary tissues could be assigned to one of four developmental categories: unemerged buds, emerged buds, unbranched lateral branches or secondarily branched lateral branches. In -R plants, branch development was retarded, with the retardation becoming increasingly pronounced as the number of -R phytomers on the primary stolon increased. Retarded elongation of the internodes of lateral shoots on -R plants resulted in the formation of a distinct fifth developmental category: short shoots (defined as branches with two or more leaves but with mean internode length equal to, or less than, 10% of that of the immediately proximal internode on the parent stolon) which had reduced phytomer appearance rates but retained the potential to develop into lateral branches. Transfer of +R plants to -R conditions, and vice versa, after 66 d demonstrated that subsequent branch development was wholly under the control of the youngest nodal root present, regardless of the age and number of root systems proximal to it.

Branching responses of a plagiotropic clonal herb to localised incidence of light simulating that reflected from vegetation.

Plants sense the presence of neighbouring vegetation through phytochrome photoreceptors perceiving a lowered red to far-red ratio (R:FR) of light reflected from such vegetation. We hypothesised that it would be advantageous for the grassland clonal herb, Trifolium repens, to have an inhibitory branching response to perception by leaves of light reflected from neighbouring vegetation (i.e. light with lowered R:FR ratio) but have no response to interception of such light by the plagiotropic stem. We tested whether photoreception of reflected light by plagiotropic stems resulted in a different branching response to photoreception by leaves and whether leaf ontogeny influenced the response. To simulate light reflected from vegetation, FR light-emitting-diodes were used to supplement controlled environment room light so that the R:FR ratio, but not the photosynthetic photon fluence rate, of light incident at the stem or leaf of a phytomer of T. repens was lowered from 1.20 to 0.25. The plagiotropic stems were unresponsive to light simulating that reflected from vegetation. This response differs from that of stems of orthotropic species, indicating that plagiotropic stems have evolved an organ-specific photobiology. Treatment of the mature leaf with light of lowered FR ratio reduced phytomer production only of the branch in the axil of the treated leaf. Similar treatment of the immature leaf retarded, in addition, branching at basal phytomers on the same side of the primary stem axis. Thus the response to light simulating that reflected from neighbouring vegetation depended upon whether the light was incident at the stem or the leaf and on the stage of leaf development. We argue that such responses improve the performance and fitness of T. repens within grassland habitats by allowing axillary buds on plagiotropic stems to branch freely when stems are in receipt of light reflected from vegetation while leaves are in full light.

Effect of light quality (red: far-red ratio) and defoliation treatments applied at a single phytomer on axillary bud outgrowth in Trifolium repens L.

We studied the effects of light quality and defoliation on the rate of phytomer appearance and axillary bud outgrowth in white clover. The treatments were applied to one phytomer, a phytomer being defined as the structural unit comprising a node, internode, axillary bud, subtending leaf and two nodal root primordia. Light of a low red:far-red (R:FR) ratio (0.27) was applied to a "target" phytomer either (i) within the apical bud and then to the axillary bud after emergence of the phytomer from the apical bud, or (ii) to the axillary bud only after emergence. The light conditions were directed to these specific parts of the plant by collimating light from small FR light-emitting diodes; with this technique we were able to change the light quality without any change in the level of photosynthetically active radiation. The subtending leaf of the target phytomer was retained or defoliated when it had emerged from the apical bud. FR light applied from the time the phytomer was within the apical bud caused a delay in branch appearance at the target phytomer. In contrast, direct treatment of the axillary bud with FR light after it had emerged from the apical bud did not result in any delay in branch appearance. As the light treatment of the apical bud may have changed the light environment of any of the organs contained in the bud we were unable to ascribe the delay in branch appearance to light perception by any particular organ. However, indirect evidence leads to the conclusion that the likely site of light perception was the developing leaf subtending the axillary bud while it was the outermost phytomer within the apical bud. These results do not support the hypothesis that the R:FR ratio of light incident at an axillary bud site is the environmental factor that controls bud development. Defoliation of the unfolding leaf reduced the rate of phytomer appearance on the main stolon but had no immediate effect on branch appearance. As a consequence there was a reduction in the number of phytomers between the stolon apical meristem and the first phytomer with a branch. This is frequently taken to indicate a relaxation of apical dominance, but in this case was found not to involve a direct effect on bud activity. A current model of white clover growth suggests that there is integration of activity between apical meristems but independence of activity and response to the local micro-environment by axillary buds. In contrast, we found that (i) defoliation reduced phytomer appearance only at the main stolon apical meristem and not at all the meristems in the plant and (ii) that a change in the local light environment of an axillary bud had no discernible effect on bud activity once the bud had emerged from the apical bud but could delay branching if applied before emergence. These results are at variance with the predictions of the model.