Effects of fertiliser addition and subsequent gopher disturbance on a serpentine annual grassland community

Summary Application of slow release fertiliser to small (0.5x1 m) plots within a serpentine annual grassland community led to significant increases in above-ground biomass and a shift in species relative abundances. In fertilised plots the native forb species which usually dominate the grassland were almost totally replaced by grasses. In the years following initial fertiliser application, a heavy mulch formed from the previous year's grass growth allowed establishment of grass species such as Bromus mollis but significantly reduced forb establishment. Gopher disturbance of fertilised plots in the second and third years of the experiment effectively removed the grass mulch and allowed re-establishment of forb species.     

Plant and soil nitrogen dynamics in California annual grassland

Seasonal changes in soil water and nitrogen availability were related to the phenology and growth of plants in California annual grassland. Plant accumulation of nitrogen was mainly confined to two short periods of the year: fall and early spring. At these times, plants were in the vegetative growth phase, roots were growing rapidly and soil moisture was high. During these periods, soil nitrate was low or depleted. High flux of nitrogen in this ecosystem, however, is indicated by the rapid disappearance of the previous year's detrital material, high microbial biomass, and high mineralizable nitrogen and nitrification potential.At the end of the summer drought, significant amounts of the previous year's detrital material had disappeared, chloroform-labile N (expressed as microbial biomass N) was at its seasonal maximum, and soil inorganic nitrogen pools were high. This suggests inorganic nitrogen flux during the drought period. The drought escaper life history characteristics of annual grasses in California annual grassland, however, may prevent plants from utilizing available nitrogen during a large part of the year.     

The mean depth of soil water uptake by two temperate grassland species over time subjected to mild soil water deficit and competitive association

Little is known concerning the soil water use dynamics of white clover (WC) and ryegrass (RG) grown in mixtures. A greenhouse study, on a deep soil, was conducted to determine the mean depth of soil water uptake of WC and RG plants grown in a competitive association and subjected to a moderate soil water deficit. Plant growth period simulated that experienced by newly sown grassland in temperate regions. Three irrigation solutions, each containing a different hydrogen isotope (deuterium) concentration, expressed as delta notation (), were provided at three different soil depths through specially constructed tubes and containers (0.50 m diameter, 1 m depth) in order to create a soil deuterium profile gradient. Young leaves and not the entire plant were harvested in order to preserve the competitive plant association over time. Patterns of leaf D value were constant for both WC and RG. Lower leaf D values in RG compared to WC was attributed to RG more efficient stomatal control. Increases in the mean depth of soil water uptake as soil water deficit increased was similar between plants. The mean depth of soil water uptake of WC was at all times greater than that of RG. After 3 months of competitive growth, WC roots obtained water from a soil depth 30% greater than that of RG. In our experimental conditions, the ability of WC to obtain water from substantially lower soil depths may give it a competitive advantage over RG during the period subsequent to pasture sowing if surface soil water deficits are experienced and deeper soil layers contain water.     

Plant and soil nitrogen dynamics in mediterranean grasslands: a comparison of annual and perennial grasses

Summary The predominance of annual species in the rangelands of southwestern Spain is not due only to climatic factors but is also strongly influenced by grazing management. Manipulating the grazing system in an experimental plot gave a vegetation structure with patches of annual grasses (mainly Vulpia ssp. and Bromus hordeaceus) and patches of perennial grasses (mainly Phalaris aquatica). This vegetation change allowed us to test the hypothesis that life-cycle differences between annual and perennial grasses affect soil nitrogen availability and plant uptake. Nitrogen availability, measured by in situ incubation, and nitrogen uptake were measured through the growing period (October to June). Amounts of in situ mineralized nitrogen over the whole growth phase were more important for soils supporting perennials (37 ppm) than for soils supporting annuals (27 ppm). The difference between the mineral nitrogen produced in situ and the mineral nitrogen accumulated during the same time in the soil allowed an estimation of the maximum mineral nitrogen quantity which can be taken up by the vegetation during each incubation period. The quantities accumulated over the year were 47 and 38 ppm (or 103 and 83 kg/ha) for soils supporting perennials and annuals respectively. For the same period, amounts of nitrogen immobilized in biomass production were 90 and 70 kg/ha for perennials and annuals respectively. During the autumn, a large proportion of mineral nitrogen was leached from soils supporting annual plants which had only just commenced germination. By contrast, the ability to use mineral nitrogen as soon as autumn rains occurred gave a competitive advantage to the perennial species, but only if they were protected from grazing during this period. The higher mineralization and use of this nitrogen reserve by perennials indicates that they made more efficient use of nitrogen resources than annuals, and validate the initial hypothesis.     

Effect of gibberellic acid on ion uptake selectivity in pea seedlings

Gibberellic acid reduced the uptake of nitrogen and phosphorus relative to the cations, a common reponse in the three pea cultivars studied. In addition, in the cv. Progress, it increased the uptake of calcium relative to magnesium and potassium. No effect in the proportion in which cations are absorbed was noticeable in the other two varieties. Ion uptake is modified by gibberellic acid through its influence on the sink strength of the shoot, the size and geometry of the root system, and the selectivity in uptake. The overall effect may result in a stimulation or an inhibition, depending on the ion considered and the pea cultivar.Abbreviation GA₃ gibberellic acid     

Plant species-specific changes in root-inhabiting fungi in a California annual grassland: responses to elevated CO2 and nutrients

Five co-occurring plant species from an annual mediterranean grassland were grown in monoculture for 4 months in pots inside open-top chambers at the Jasper Ridge Biological Preserve (San Mateo County, California). The plants were exposed to elevated atmospheric CO₂ and soil nutrient enrichment in a complete factorial experiment. The response of root-inhabiting non-mycorrhizal and arbuscular mycorrhizal fungi to the altered resource base depended strongly on the plant species. Elevated CO₂ and fertilization altered the ratio of non-mycorrhizal to mycorrhizal fungal colonization for some plant species, but not for others. Percent root infection by non-mycorrhizal fungi increased by over 500% for Linanthus parviflorus in elevated CO₂, but decreased by over 80% for Bromus hordeaceus. By contrast, the mean percent infection by mycorrhizal fungi increased in response to elevated CO₂ for all species, but significantly only for Avena barbata and B. hordeaceus. Percent infection by mycorrhizal fungi increased, decreased, or remained unchanged for different plant hosts in response to fertilization. There was evidence of a strong interaction between the two treatments for some plant species and non-mycorrhizal and mycorrhizal fungi. This study demonstrated plant species- and soil fertility-dependent shifts in below-ground plant resource allocation to different morpho-groups of fungal symbionts. This may have consequences for plant community responses to elevated CO₂ in this California grassland ecosystem. Received: 2 June 1997 / Accepted: 22 August 1997     

The persistence of calcareous grassland species in the soil seed bank under developing and established scrub

The relationships between the composition of the soil seed bank, the field layer vegetation, and the scrub canopy were investigated along a 69 m transect, grading from incipient woodland, through scrub, into intensively rabbit-grazed calcareous grassland. The results are used to assess the persistence of species associated with open calcareous grassland in the seed bank under developing scrub. Scrub age, composition and density, changed along the transect from the woodland to open grassland. A total of 35 forb and grass species were found in the field layer. The pattern evident in the scrub layer was also reflected in the herbaceous vegetation. The field layer in the most closed portion of the transect, where the scrub was oldest, was dominated by shade-tolerant species normally associated with woodland habitats. The abundance of these species decreased along the transect as the scrub age declined, and the field layer became increasingly dominated by species typical of open grassland. A total of 47 species germinated from the seed bank. Few species were recorded in the seed bank along the entire length of the transect. Overall, the seed bank was dominated by Hypericum perforatum and Centaurium erythraea, which accounted for 38.2% and 28.6% of emerging seedlings respectively. As with a number of similar studies, the composition of the seed bank had a low correspondence with the composition of the field layer vegetation. The results also emphasise that the composition of the seed bank can be viewed as an ecological palimpsest, with germinable seed of species from each stage of the old-field succession occurring in the soil. The seed bank is an important component in the re-vegetation of an area after disturbance such as scrub removal. This study supports the findings of previous research in showing that relatively few characteristic calcareous grassland species form persistent seed banks. The soil seed bank would therefore appear to be of limited value in the restoration of such grassland following scrub removal.     

Effect of climate on the growth of annual rings in the main roots of perennial forbs in an Inner Mongolian semi‐arid grassland,China

Question: Is there a pattern in growth of annual rings in roots of perennial forbs in relation to climate and climate extremes in grassland ecosystems? Location: Semi‐arid grassland in Duolun (42°27′N, 116°41′E, 1380 m a.s.l.), central Inner Mongolia, China. Methods: Main roots of three perennial species, Potentilla anserina L., Cymbaria dahurica L. and Lespedeza daurica Schindl., were sampled. Cross‐sections (10–15‐μm thick) were produced from the proximal end of sampled roots using a sledge microtome. Annual growth rings in the main roots were identified and measured by differentiating between earlywood and latewood in the secondary xylem. Relationships between annual growth rings and monthly mean temperature and total monthly precipitation were identified using correlation analysis. Differences in an annual ring width to the previous and following years were examined by calculating a distinctness score. Results: The three perennial forbs showed clearly demarcated annual growth rings in all individuals and the same fluctuation patterns. Their ring widths were generally positively correlated with precipitation from April to October (except for August) and with temperature from February to June (except June for L. daurica), September to October, and the annual mean. Strong deviations of annual ring widths from their neighbour rings were observed in 1998 and 2000. The trend of absolute distinctness scores (D_m) increased significantly from 1988 to 2003, indicating an increase in the frequency of annual ring width variation. Conclusions: Annual growth rings in the main roots of three perennial forb species can be used as an indicator of the influence of climate on below‐ground grassland growth. The change in below‐ground conditions and effects on the functioning of grassland should receive more attention in future studies.     

Preferential uptake of soil nitrogen forms by grassland plant species

In this study, we assessed whether a range of temperate grassland species showed preferential uptake for different chemical forms of N, including inorganic N and a range of amino acids that commonly occur in temperate grassland soil. Preferential uptake of dual-labelled (¹³C and ¹⁵N) glycine, serine, arginine and phenylalanine, as compared to inorganic N, was tested using plants growing in pots with natural field soil. We selected five grass species representing a gradient from fertilised, productive pastures to extensive, low productivity pastures (Lolium perenne, Holcus lanatus, Anthoxanthum odoratum, Deschampsia flexuosa, and Nardus stricta). Our data show that all grass species were able to take up directly a diversity of soil amino acids of varying complexity. Moreover, we present evidence of marked inter-species differences in preferential use of chemical forms of N of varying complexity. L. perenne was relatively more effective at using inorganic N and glycine compared to the most complex amino acid phenylalanine, whereas N. stricta showed a significant preference for serine over inorganic N. Total plant N acquisition, measured as root and shoot concentration of labelled compounds, also revealed pronounced inter-species differences which were related to plant growth rate: plants with higher biomass production were found to take up more inorganic N. Our findings indicate that species-specific differences in direct uptake of different N forms combined with total N acquisition could explain changes in competitive dominance of grass species in grasslands of differing fertility.     

The effects of elevated atmospheric CO2 on the amount and depth distribution of plant water uptake in a California annual grassland

Soil moisture profiles can affect species composition and ecosystem processes, but the effects of increased concentrations of atmospheric carbon dioxide ([CO₂]) on the vertical distribution of plant water uptake have not been studied. Because plant species composition affects soil moisture profiles, and is likely to shift under elevated [CO₂], it is also important to test whether the indirect effects of [CO₂] on soil water content may depend on species composition. We examined the effects of elevated [CO₂] and species composition on soil moisture profiles in an annual grassland of California. We grew monocultures and a mixture of Avena barbata and Hemizonia congesta– the dominant species of two phenological groups – in microcosms exposed to ambient (～370 μmol mol^?1) and elevated (～700 μmol mol^?1) [CO₂]. Both species increased intrinsic and yield‐based water use efficiency under elevated [CO₂], but soil moisture increased only in communities with A. barbata, the dominant early‐season annual grass. In A. barbata monocultures, the [CO₂] treatment did not affect the depth distribution of soil water loss. In contrast to communities with A. barbata, monocultures of H. congesta, a late‐season annual forb, did not conserve water under elevated [CO₂], reflecting the increased growth of these plants. In late spring, elevated [CO₂] also increased the efficiency of deep roots in H. congesta monocultures. Under ambient [CO₂], roots below 60 cm accounted for 22% of total root biomass and were associated with 9% of total water loss, whereas in elevated [CO₂], 16% of total belowground biomass was associated with 34% of total water loss. Both soil moisture and isotope data showed that H. congesta monocultures grown under elevated [CO₂] began extracting water from deep soils 2 weeks earlier than plants in ambient [CO₂].     

Nitrite in the root zone and its effects on ion uptake and growth of wheat seedlings

The immediate and posteffects of various concentrations of NaNO₂ on ion uptake of wheat ( Triticum aestivum L. cv. GK Öthalom) seedlings were studied at different pH values. Without pretreatment, the higher the concentration of NaNO₂ the greater was the decrease in uptake of K⁺ into the roots, both at pH 4 and pH 6. At pH 6 but not at pH 4 the reverse was true when the seedlings were pretreated with NaNO₂. Due to the high Na⁺ content of the roots, an effect of Na⁺ in this process cannot be excluded. Nitrite was taken up by the roots more rapidly than nitrate. Nitrite at 0.1 m M in the medium induced the development of an uptake system for both NO⁻₂ and NO⁻₃ in wheat roots. At higher concentrations pretreatment with NO⁻₂ decreased NO⁻₃ uptake by the roots, but NO₃ did not inhibit the uptake of NO₂. The toxic effect of NO⁻₂ was strongly pH dependent. Lower pH of the external solution led to an increased inhibition by NO⁻₂ of both ion uptake and growth of seedlings. The inhibitory effect of NO⁻₂ differed considerably for roots and shoots. The roots and especially the root hairs were particularly sensitive to NO⁻₂ treatment.     

Effects of common soil anions and pH on the uptake and accumulation of perchlorate in lettuce

A mechanistic understanding of perchlorate () entry into plants is important for establishing the human health risk associated with consumption of contaminated produce and for assessing the effectiveness of phytoremediation. To determine whether common soil anions affect uptake and accumulation in higher plants, a series of competition experiments using lettuce (Lactuca sativa L.) were conducted between (50 nM) and (4–12 mM), (1–10 mM), or Cl⁻ (5–15 mM) in hydroponic solution. The effects of (0–5 mM) and pH (5.5–7.5) on uptake were also examined. Increasing in solution significantly reduced the amount of taken up by green leaf, butter head, and crisphead lettuces. Sulfate and Cl⁻ had no significant effects on uptake in lettuce over the concentrations tested. Increasing pH significantly reduced the amount of taken up by crisphead and green leaf lettuces, whereas increasing significantly reduced uptake in butter head lettuce. The inhibition by across all lettuce genotypes suggests that may share an ion carrier with , and the decrease in uptake with increasing pH or provides macroscopic evidence for cotransport across the plasma membrane.     

Glucose uptake and phosphorylating activities in two species of slow-growing Rhizobium

Abstract Glucose uptake and phosphorylating activities were studied in two strains of slow-growing Rhizobium: Rhizobium japonicum (USDAI-110) and cowpea Rhizobium (USDA3278). Cultured cells of both species actively took up glucose, and at least two systems appeared to be involved, whereas purified bacteroids of both species failed to accumulate glucose actively. In both cell types, glucose phosphorylation was ATP-dependent, and no evidence was obtained for a phosphoenolpyruvate-dependent glucose phosphotransferase system.     

Effect of soil compaction on root growth and uptake of phosphorus

Summary Zea mays L. andLolium rigidum Gaud. were grown for 18 and 33 days respectively in pots containing three layers of soil each weighing 1 kg. The top and bottom layers were 100 mm deep and they had a bulk density of 1200 kg m^–3, while the central layer of soil was compacted to one of 12 bulk densities between 1200 and 1750 kg m^–3. The soil was labelled with³²P and³³P so that the contribution of the different layers of soil to the phosphorus content of the plant tops could be determined. Soil water potential was maintained between –20 and –100 kPa.Total dry weight of the plant tops and total root length were slightly affected by compaction of the soil, but root distribution was greatly altered. Compaction decreased root length in the compacted soil but increased root length in the overlying soil. Where bulk density was 1550 kg m^–3, root length in the compacted soil was about 0.5 of the maximum. At that density, the penetrometer resistance of the soil was 1.25 and 5.0 MPa and air porosity was 0.05 and 0.14 at water potentials of –20 and –100 kPa respectively, and daytime oxygen concentrations in the soil atmosphere at time of harvest were about 0.1 m³m^–3. Roots failed to grow completely through the compacted layer of soil at bulk densities 1550 kg m^–3. No differences were detected in the abilities of the two species to penetrate compacted soil.Ryegrass absorbed about twice as much phosphorus from uncompacted soil per unit length of root as did maize. Uptake of phosphorus from each layer of soil was related to the length of root in that layer, but differences in uptake between layers existed. Phosphorus uptake per unit length of root was higher from compacted than from uncompacted soil, particularly in the case of ryegrass at bulk densities of 1300–1500 kg m^–3.     

Effect of soil drying on growth,biomass allocation and leaf gas exchange of two annual grass species

Influence of short-term water stress on plant growth and leaf gas exchange was studied simultaneously in a growth chamber experiment using two annual grass species differing in photosynthetic pathway type, plant architecture and phenology:Triticum aestivum L. cv. Katya-A-1 (C₃, a drought resistant wheat cultivar of erect growth) andTragus racemosus (L.) All. (C₄, a prostrate weed of warm semiarid areas). At the leaf level, gas exchange rates declined with decreasing soil water potential for both species in such a way that instantaneous photosynthetic water use efficiency (PWUE, mmol CO₂ assimilated per mol H₂O transpired) increased. At adequate water supply, the C₄ grass showed much lower stomatal conductance and higher PWUE than the C₃ species, but this difference disappeared at severe water stress when leaf gas exchange rates were similarly reduced for both species. However, by using soil water more sparingly, the C₄ species was able to assimilate under non-stressful conditions for a longer time than the C₃ wheat did. At the whole-plant level, decreasing water availability substantially reduced the relative growth rate (RGR) ofT. aestivum, while biomass partitioning changed in favour of root growth, so that the plant could exploit the limiting water resource more efficiently. The change in partitioning preceded the overall reduction of RGR and it was associated with increased biomass allocation to roots and less to leaves, as well as with a decrease in specific leaf area. Water saving byT. racemosus sufficiently postponed water stress effects on plant growth occurring only as a moderate reduction in leaf area enlargement. For unstressed vegetative plants, relative growth rate of the C₄ T. racemosus was only slightly higher than that of the C₃ T. aestivum, though it was achieved at a much lower water cost. The lack of difference in RGR was probably due to growth conditions being relatively suboptimal for the C₄ plant and also to a relatively large investment in stem tissues by the C₄ T. racemosus. Only 10% of the plant biomass was allocated to roots in the C₄ species while this was more than 30% for the C₃ wheat cultivar. These results emphasize the importance of water saving and high WUE of C₄ plants in maintaining growth under moderate water stress in comparison with C₃ species.     

Plant respiration in relation to growth, maintenance, ion uptake and nitrogen assimilation

Abstract Respiration in plants is generally observed to comprise two components: one proportional to the growth rate and the other to the plant dry mass. These components are usually interpreted as being related to the growth of new plant material and maintenance of existing plant material, respectively. By analysing data in this way, the respiratory costs of both structural synthesis and maintenance are observed to be greater in the root than the shoot. This contradicts current understanding of the biochemistry of the processes involved. The basic model is developed to incorporate three additional processes. The first is the cost of ion uptake for plant growth. The second allows for the fact that the site of nitrogen assimilation into amino acids may differ from the site of utilization for protein synthesis: when ammonium is supplied, this is incorporated immediately into amino acids owing to its toxicity to the plants; when nitrate is supplied it may be reduced either in the shoot or root, or both, and subsequently transported around the plant for utilization. The third process to be included is an energy cost for the uptake of ions to balance efflux from the root system. The theory is consistent with experimental observation and provides a means of understanding and interpreting respiration and nitrogen metabolism in plants.