Carol A. Marriott

Spatial variability of soil total C and N and their stable isotopes in an upland Scottish grassland

As preparation for a below ground food web study, the spatial variability of three soil properties (total N, total C and pH) and two stable isotopes (δ13C and δ15N of whole soil) were quantified using geostatistical approaches in upland pastures under contrasting management regimes (grazed, fertilised and ungrazed, unfertilised) in Scotland. This is the first such study of upland, north maritime grasslands. The resulting patterns of variability suggest that to obtain statistically independent samples in this system, a sampling distance of ≥13.5 m is required. Additionally, temporal change (a decline of 1‰) was observed in whole soil δ15N for the grazed, fertilised plot. This may have been caused by new inputs of symbiotically-fixed atmospheric N2.

Stable isotope natural abundances of soil, plants and soil invertebrates in an upland pasture

Abstract In an exploratory study of below-ground trophic relations, natural abundances of the stable isotope pairs 13C/12C and 15N/14N (δ13C and δ15N) were measured on samples of plant shoots, whole soil and soil invertebrates taken in 1994 from two contrasting treatments of a pre-existing experiment: (1) continued grazing by sheep, with N:P:K fertiliser additions from 1990 onward; and (2) no added fertiliser, but sheep grazing removed entirely. Stepwise trophic increases were documented better by seasonal averages of δ13C and δ15N and by seasonal trends, composed of data collected on several occasions, than by instantaneous values. Seasonal changes in plant monocot vs dicot differences for shoot δ13C and δ15N were detected from patterns over several individual sampling dates; instantaneous samples were neither statistically significant nor qualitatively interpretable. Significant isotopic differences between treatments were evident in invertebrates only as seasonal averages or trends. Seasonal variations of δ13C and δ15N in earthworms and slugs may reflect previously unsuspected invertebrate behaviour. Whole soil δ13C was static through time and across treatments. Whole soil δ15N changed seasonally, an effect consistent with 15N/14N fractionation, e.g. during denitrification.

Transfer of legume nitrogen to associated grass

Abstract In improved hill-peat a rapid increase in rhizosphere soil bacteria occurs following clover defoliation. The increase in bacterial-cell N is equivalent to 19% of the N released from the clover roots in the first 5 days after cutting. Subsequent ammonification of the immobilized N is dependent on the resistance of the microbial fraction to further degradation. In three fertilized hill soils examined, [15N]labelled bacterial N was mineralized quite rapidly, some 20% of the bacterial N being taken up by grass over 17 days. Under glasshouse conditions the rate of mineralization is different in the three soils examined, the peat providing a more favourable environment than brown earth for bacterial N transfer.

The effect of sheep urine on clover performance in a grazed upland sward

The effect of artificially applied urine on clover performance in a perennial ryegrasswhite clover sward grazed by sheep was examined during summer. Sheep urine or deionized water (51/m 2 ) was applied to areas (2·70 × 1·25 m) which were protected from subsequent excretal return by graze-through cages. Grass and clover populations, nitrogen-fixing activity, soil nitrogen and soil pH were monitored over the following 90-day period. Urine reduced clover population density, stolon length and dry weight but had little effect on number of grass tillers. Nitrogen-fixing activity of clover was reduced initially to less than 30% of control values, perhaps owing to high levels of soil inorganic nitrogen inhibiting fixation, although osmotic effects due to the high salt concentration in urine cannot be discounted. The lower levels of activity at later dates were due largely to reduced clover in urine-treated areas. Soil pH (0–5 cm) was increased by about 0–5 units 3 days after urine application, with a smaller increase in the 5–10 cm horizon. Recovery of urine nitrogen in the soil mineral N pool was at best 27% of the added nitrogen. The results are discussed in terms of the significance of urine-affected areas in determining a patchy distribution of clover in grazed swards; the role of animals in the transfer of fixed N from clover to grass is considered.

Above-ground grazing affects floristic composition and modifies soil trophic interactions

Abstract There are few data on the functional inter-relationships between above- and below-ground components of soil ecosystems. Here, we report changes in below-ground soil invertebrate trophic relationships (manifested as alterations in stable isotope natural abundances, δ 13 C and δ 15 N) that arose in association with the removal of sheep grazing and from the resulting changes in above-ground floristic composition. Consequent to grazing removal, Lolium perenne L. (perennial rye-grass) was replaced as the dominant plant species in ungrazed treatments by Ranunculus repens L. (creeping buttercup), a species with more 13 C-enriched foliage. Consequently, all invertebrate functional groups studied, but not whole soil, were more 13 C-enriched in ungrazed treatments. Earthworms (detritivore) from grazed treatments were significantly 15 N-enriched compared with earthworms from ungrazed treatments. In contrast, slug (herbivore) δ 15 N exhibited no treatment effect. Reasons for this are unclear but may be related to the effects of above-ground grazing on the composition of below-ground microbial/microfaunal communities. Omnivores/carnivores (beetles and spiders), were more 15 N-enriched than primary producers in the grazed than in the ungrazed treatments (6 vs. 4‰) suggesting a longer below-ground foodchain in the grazed plots. The cessation of fertilizer application had no comparable effects on below-ground trophic relationships.

Seasonal dynamics of leaf extension and losses to senescence and herbivory in extensively managed sown ryegrass–white clover swards

Tissue flow measurements of leaf material in Lolium perenne (perennial ryegrass), Agrostis capillaris or Poa annua , and Trifolium repens (white clover) were made at three upland sites in Scotland (Hartwood, Sourhope and Fasque) in 1992/93 to determine if there were differences in seasonal growth, senescence and losses to herbivory between species in their response to more extensive sward management. The measurements were made monthly from May until October in established predominantly perennial ryegrass/white clover swards receiving three different treatments. One treatment received a conventional annual fertilizer application of 140 kg N/ha plus maintenance P and K and was grazed by sheep at a sward height of 4 cm (4F), whereas the other two treatments were unfertilized and grazed to maintain a sward height of 4 cm (4U) or 8 cm (8U). Significant sources of the variation in leaf appearance, increase in green lamina/petiole length (leaf extension), senescence and losses to herbivory were attributable to site, sward, species and date of measurement. The rate of leaf extension for all three measured species was less in 4U than 4F swards, and less in 4U than 8U swards. Leaf extension of L. perenne exceeded that of the other species, even in unfertilized swards, but rates of leaf appearance were less. There was some evidence in spring of a reduction in net growth as a consequence of removing fertilizer inputs and maintaining a sward height of 4 cm. Species differences in the losses of leaf tissue to herbivory were dependent on sward management. In the 4F treatment, leaf loss to herbivory from L. perenne tillers was greater than that from either A. capillaris , P. annua or T. repens in May, June and September. In the 4U treatment more leaf tissue was also lost from L. perenne than from T. repens . In contrast, there was no difference between grass species in losses to herbivory in either unfertilized sward. The responses of species to changes in fertilizer and grazing management were similar at three sites of differing fertility. The results are discussed in relation to plant competition and species dynamics in extensively managed swards.

Senescence and decomposition of white clover stolons in grazed upland grass/clover swards

The fate of marked sections of stolons of white clover (Trifolium repens) over a 50-week period from May 1987 was followed in grazed grass/clover swards maintained at 5-cm sward surface height with and without N fertiliser. There was little effect of N treatment on the pattern of survival of stolon sections. The proportion of live stolons recovered decreased during the experiment, and in May 1988 on average only 29% of the marked sections remained alive. At all harvests only a small percentage of stolon sections showed signs of senescence; the maximum percentage, on average 20% of those marked, occurred in autumn, 15–20 weeks after marking. Following this period, i.e. in late autumn/winter, the most rapid increase in percentage of decomposed stolons was measured. Over 50% of stolon sections were buried within the 5-week period following marking and nearly all were buried after 20 weeks; generally a much smaller proportion of stolon tips was buried. Nutrient concentrations of N, P and K fell to their lowest levels in autumn, before increasing in the following spring. Results are discussed in relation to the cycling of nutrients via stolon senescence.

Does the uniform packing of sand in a cylinder provide a uniform penetration resistance? A method for screening plants for responses to soil mechanical impedance

Sand packed to a constant dry bulk density, is frequently used as an artificial growth medium in which to simulate the effects of a constant mechanical impedance on root growth. This research aimed to determine whether conventional packing resulted in constant mechanical impedance and to test alternative packing regimes. Perspex cylinders 300 mm tall with a 49 mm internal diameter were packed with moist sand to uniform and varying bulk densities to examine which type of packing gave the greatest uniformity of penetration resistance (PR) with depth. The cylinders packed to a constant bulk density (1.48, 1.55, and 1.6 Mg m-3) all had measured PR profiles which increased markedly with depth by approximately 1, 1.5 and 3 MPa, respectively, within the top 100 mm. Between 100–300 mm depth, these same cylinders showed reductions in PR of up to 1, 2 and 2 MPa respectively. These results show that sand packed to a constant bulk density with depth would not provide a uniform mechanical impedance to plant roots.By packing sand to different bulk densities at different depths, we obtained packed cylinders that had much more uniform PR profiles (with average values of 0.25, 1.40 and 2.30 MPa). Below a depth of 50 mm, the coefficients of variation for replicate cylinders packed in this way were 12%, 5% and 18% for the 0.25, 1.40, and 2.30 MPa treatments respectively. For experiments with single plants, the lower PR values that were unavoidable near to the surface (< 50 mm) can be avoided by sowing seeds at the base of a funnel inserted into the cylinder. Treatments such as these can provide reproducible growth media, with adequate water/nutrient and aeration status for the study of plant response to uniform mechanical impedance.

The effect of temperature and nitrogen interactions on growth and nitrogen assimilation of white clover

White clover plants were grown for 97 days under two temperature regimes (20/15°C and 8/5°C day/night temperatures) and were supplied with either small amounts (a total of 80 mg N pot−1) of ammonium (NH4+) or nitrate (NO3−) nitrogen, or received no mineral N and relied on N2 fixation. Greatest growth and total leaf area of clover plants occurred in N2 fixing and NO3−-fed plants grown at 20/15°C and poorest growth occurred in NH4+-fed plants grown at 8/5°C. Nodule mass per plant was greater at 8/5°C due to increased nodule numbers rather than increased dry weight per nodule. This compensated to some extent for the reduced N2-fixing activity per unit dry weight of nodule tissue found at the low growth temperature up to 116 d after sowing, but thereafter both activity per nodule dry weight and activity per plant were greater at the low temperature. Highest nitrate reductase activity (NRA) per g fresh weight and total activity per leaf, petiole or root occurred in NO3−-fed plants at 8/5°C. Low growth temperature resulted in a greater partitioning of total plant NRA to the roots of NO3−-fed plants. The results are considered in relation to the use of N fertiliser in the spring under field conditions.