Photosynthetic responses of Larrea tridentata to a step-increase in atmospheric CO2at the Nevada Desert FACE Facility

Of all terrestrial ecosystems, the productivity of deserts has been suggested to be the most responsive to increasing atmospheric CO₂. The extent to which this prediction holds will depend in part on plant responses to elevated CO₂under the highly variable conditions characteristic of arid regions. The photosynthetic responses ofLarrea tridentata , an evergreen shrub, to a step-increase in atmospheric CO₂(to 550 μmolmol⁻¹) were examined in the field using Free-Air CO₂Enrichment (FACE) under seasonally varying moisture conditions. Elevated CO₂substantially increased net assimilation rate (A_net) in Larrea during both moist and dry periods of the potential growing season, while stomatal conductance (g_s) did not differ between elevated and ambient CO₂treatments. Seasonal and diurnal gas exchange dynamics in elevated CO₂mirrored patterns in ambient CO₂, indicating that elevated CO₂did not extend photosynthetic activity longer into the dry season or during more stressful times of the day. Net assimilation vs. internal CO₂(A/C_i) responses showed no evidence of photosynthetic down-regulation during the dry season. In contrast, after significant autumn rains, A_max(the CO₂saturated rate of photosynthesis) and CE (carboxylation efficiency) were lower in Larrea under elevated CO₂. In situ chlorophyll fluorescence estimation ofLarrea Photosystem II efficiency (F_v/F_m) responded more to water limitation than to elevated CO₂. These findings suggest that predictions regarding desert plant responses to elevated CO₂should account for seasonal patterns of photosynthetic regulatory responses, which may vary across species and plant functional types.     

Photosynthetic response to different light intensities, water status and leaf age of two Berberis species (Berberidaceae) of Patagonian steppe, Argentina

Net photosynthetic rate (A) and stomatal conductance (gs) from leaves of Berberis buxifolia and Berberis heterophylla saplings were measured under different conditions of radiation and water availability and for leaves of different ages. Comparative studies of basic physiological performance would give insight how these shrubs survive and grow under this austral extreme ecosystem. B. buxifolia had higher A values than B. heterophylla for all light intensities measured. Values of gs for both Berberis species also declined as light intensity decreased. There was a negative relationship between A and water stress expressed as pre-dawn leaf water potential (ψ_lp) where the reduction in gs was the main factor that reduced A in both species. Also, A and gs of Berberis plants increased rapidly during the first days after leaf unfolding, reached a maximum value when the leaf was completely expanded, and then declined. These results could be used to estimate the responses of saplings to environmental conditions under the dynamic of natural stands in this austral extreme ecosystem and for restoration plans.     

Quantitative evaluation of the counterbalance between photosynthetic stimulation and depression caused by low partial pressure of O2 and CO2 in alpine atmospheres

To evaluate the effects of low atmospheric pressure on leaf photosynthesis, we compared the photosynthesis of identical leaves of Fagus crenata at lowland (0 m a.s.l.) and at highland (2360 m a.s.l.). At the high altitude, the atmospheric pressure and partial pressure of CO₂ at intercellular air spaces in the leaf (C_i360) decreased to 77% and 78% of those at the low altitude, respectively. On the other hand, the efficiency of photosynthetic CO₂-utilization was apparently higher at the high altitude because of a mitigation of the O₂-inhibition of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) under low ambient partial pressure of O₂. This stimulation of photosynthesis partly compensated a photosynthetic depression due to the low C_i360, and the net assimilation rate (A_n360) at the high altitude retained 94% of the value at the low altitude. A theoretical model indicated that the stimulation of photosynthesis at high altitudes depend on internal conductance (g_i) and/or on Rubisco content. The model demonstrated that low atmospheric pressure at high altitudes caused severe restrictions of photosynthesis when leaves had a small g_i and/or a large amount of Rubisco, whichever are repeatedly reported in alpine plants.     

Effects of drought stress on antioxidant enzyme, photosynthetic pigment and flavonoid pathway in two desert shrubs

Antioxidant enzyme activity, photosynthetic pigment content, and free malondialdehyde (MDA), as well as flavonoid content and the key enzyme activity in the flavonoid pathway were determined in two desert shrubs, Caryopteris mongolica Bunge and Reaumuria soongorica (Pall.) Maxim. under drought stress. The free MDA content was enhanced during the experimental period, which may be an indicator of oxidative stress. Superoxide dismutase (SOD), peroxidase (POD), and ascorbate peroxidase (APX) activities in C. mongholica showed a significant increase during the experiment, but catalase (CAT) activity was slightly decreased. On the other hand, POD and APX activities showed a significant increase and SOD and CAT activity data had no significant changes in R. soongorica. APX, SOD, and CAT activities were higher in R. soongorica than in C. mongholica, but MDA content was lower, indicating that the lower values of MDA were attributed to higher activities of antioxidant enzyme in R. soongorica. Chlorophyll content decreased significantly in the two shrubs during the experiment, which indicated that there was a photoprotection mechanism through reducing light absorbance by decreasing pigments content. Caretonoids content increased in C. mongholica and decreased in R. soongorica. The ratio of Chla/Chlb decreased significantly but caretonoids/Chl revealed a significant increase in the two shrubs, which could be explained as no decrease of peripheral light-harvesting complexes and a higher tolerance to drought. Total flavonoid content and the activities of phenylalanine ammonialyase (PAL) and chalcone isomerase (CHI) showed different changes between C. mongholica and R. soongorica after treatment. These values decreased in R. soongorica and increased in C. mongholica except for PAL activity. However, anthocyanin content increased in the two shrubs, indicating that there was a different regulation response in the flavonoid pathway in the two shrubs under drought stress, and anthocyanin should be an important antioxidant both in the shrubs. Our results demonstrated the different responses of antioxidant defense and drought tolerance ability between the two shrubs.     

Factors affecting the distribution, abundance and seedling survival of Mammillaria gaumeri , an endemic cactus of coastal Yucatán, México

The population status of an endemic cactus (Mammillaria gaumeri) was assessed in coastal Yucatán, México. Similar numbers of mature plants were found in open and full sun microhabitats. Plants in full sun had more ramets and were of larger diameter. There was a correlation between M. gaumeri abundance and the biomass of a disturbance indicator, Myrmecophila tibicinis (Orchidacea). Higher seedling survival in shade microhabitats and aggregated distribution of adults suggests that shrubs serve as nurse plants. Initial mortality rates of seedlings were many times higher in full sun microhabitats than in shade conditions. Herbivory had no effect on seedling survival.     

Rainfall interception by sand-stabilizing shrubs related to crown structure

On the edge of the Tengger Desert in northern China,revegetation has changed the landscape from moving dunes to stabilized dunes covered by shrubs,which further modifies the pattern of rainfall redistribution.To study rainfall interception loss by shrubs and its relationship to rainfall properties and crown structure,throughfalls passing through crowns of Artemisia ordosica Krash.and Caragana korshinskii Kom.were measured using nine PVC cups under the canopy of each of the two shrubs during 73 rain events over a three-year period,with total rainfall of 260.9 mm.Interception losses of gross rainfall by A.ordosica and C.korshinskii account for 15% and 27% of the total on a crown area basis,and 6% and 11% on a ground area basis,respectively.Individual throughfall(T) and interception(I) were significantly related to rainfall amount(Pg),duration(D),and intensity(R).Ratios of throughfall to rainfall(T/Pg) and interception to rainfall(I/Pg) were not only significantly related to Pg,D,and R,but also to shrub species,and interactions of species with crown volume(CV) and leaf area index(LAI).Under most rain events,interceptions by C.korshinskii with greater CV and LAI were significantly higher than those by A.ordosica,and more rainfall interception occurred at locations closer to the stems of the two shrubs.For C.korshinskii,I/Pg had a significant positive linear relation with CV and LAI,while T/Pg had a significant negative linear relation with them.CV has a greater influence on T/Pg and I/Pg than does LAI.Using a regression method,canopy water storage capacities are estimated to be 0.52 and 0.68 mm,and free throughfall coefficient to be 0.62 and 0.47 for A.ordosica and C.korshinskii,respectively.     

Effect of exogenous application of salicylic acid on the drought stress responses of Gardenia jasminoides

The alleviative effects of exogenous salicylic acid(SA) on plants against drought stress were assessed in Gardenia jasminoides seedlings treated with different concentrations of SA.Drought stress was simulated to a moderate level by 15% polyethylene glycol(PEG) 6000 treatment.Seedlings exposed to 15% PEG for 14 days exhibited a decrease in aboveground and underground dry mass,seedling height,root length,relative water content,photosynthetic pigment content,net photosynthetic rate(Pn),transpiration rate(Tr),stomatal conductance(Gs),and water use efficiency.In PEG-stressed plants,the levels of proline,malondialdehyde(MDA),hydrogen peroxide(H_2O_2),and electrolyte leakage rose significantly,whereas antioxidative activity,including superoxide,peroxidase,and catalase activities,declined in leaves.However,the presence of SA provided an effective method of mitigating PEG-caused physiological stresses on G.jasminoides seedlings,which depended on SA levels.PEG-treated plants exposed to SA at 0.5–1.0 mmol/L significantly eased PEG-induced growth inhibition.Application of SA,especially at concentrations of 0.5–1.0 mmol/L,considerably improved photosynthetic pigments,photosynthesis,antioxidative activity,relative water content,and proline accumulation,and decreased MDA content,H_2O_2 content,and electrolyte leakage.By contrast,the positive effects were not evident,or even more severe,in PEG+SA4 treatment.Based on these physiological and biochemical data,a suitable concentration of SA,potential growth regulators,could be applied to enhance the drought tolerance of G.jasminoides.     

灌溉量对2种灌木光合特性和水分利用效率的影响

采用野外试验,研究了灌溉量对新疆策勒绿洲木沙拐枣（Calligonum mongolicum）和花棒（Hedysarum scoparium）光合特性和水分利用效率的影响。结果表明：（1） 随着灌溉量的增加,沙拐枣和花棒的净光合速率均增加,但是蒸腾速率、气孔导度、胞间CO₂浓度没有明显增加;（2） 灌溉增加了沙拐枣的清晨水势和正午水势,并且只有灌溉和无灌溉处理的正午水势差异显著（p<0.05）;而灌溉只增加了花棒的清晨水势,并且只有充分灌溉和无灌溉的条件下差异显著（p<0.05）;（3） 在不同的灌溉条件下,沙拐枣和花棒的稳定碳同位素比率δ¹³C变化不大,沙拐枣在无灌溉、适度灌溉、充分灌溉下δ¹³C的平均值分别为-12.267、-12.147、-11.958,花棒的δ¹³C分别为-27.928、-28.068、-27.166;（4） 沙拐枣和花棒的水分利用效率均在无灌溉条件下明显高于灌溉条件下,而且一次性的灌溉不会改变其水分利用效率。     

Adaptive evolution of rbcL in Reaumuria soongarica (Tamaricaceae)

In the field of phylogenetic analyses, the rbcL gene encoded large subunit Ribulose-1,5-biphosphate carboxylase/oxygenase (Rubisco, EC4.1.1.39), which plays a crucial role in the process of photosynthesis for most terrestrial plants, has been considered to be conserved; however, recent controversy regarding rbcL conservation has appeared since it was proposed to be under natural selection within all principal lineages of land plants. In this study, by examining the variation of DNA and protein sequences among 17 species in the family Tamaricaceae, three nonsynonymous mutations were identified to be under positive selection. The favored sites were located in the alph-helix domains of Rubisco, with decreased hydrophobicity and increased entropy, which could facilitate CO₂ penetration into the active site of Rubisco. We also found that the expression level of rbcL in different genotypes of Reaumuria soongarica shifted in response to various stresses such as drought, temperature, salt, and light. This study not only sheds light on the functional/structural features of Rubisco in the evolution scenarios from C₃-like into C₄ in Tamaricaceae but also provides useful information on directing genetic performance to enhance photosynthesis efficiency of desert plants for sustaining fragile desert ecosystems; furthermore, it promotes the ability to cope with desert aridification and global warming.     

阿拉善荒漠典型植物功能群氮、磷化学计量特征

氮（N）、磷（P）作为荒漠生态系统重要的养分元素和限制性因子,在维持植物功能群组成及结构稳定、生态系统内养分循环中发挥着重要的作用。本研究沿水热梯度从东至西在阿拉善荒漠设置52个调查样地,通过对优势植物的调查取样,分析了荒漠植物不同功能（类）群N、P元素的生态化学计量特征,并就N、P含量及N：P值对荒漠植物的限制性作用进行了检验和讨论。结果显示：（1）所调查阿拉善荒漠区植物叶片的N、P含量分别为10.65±7.91 mg·g^-1和1.04±0.81 mg·g^-1,N：P值为11.53±5.06,叶片N含量与P含量及N：P值均显著正相关,P含量与N：P值显著负相关;通过对比分析认为该区植物同时受到N、P双重制约,且更易受N限制。（2）从植物生活型比较,认为草本植物叶片N、P含量均高于灌木植物叶片,而灌木植物叶片的N：P值大于草本植物;对光合作用途径分析发现,C₃植物叶片N含量高于C₄植物叶片,而C₄植物叶片P含量高于C₃植物叶片;从系统发育类型方面分析,认为单子叶植物叶片N含量低于双子叶植物叶片N含量,而P含量则高于双子叶植物叶片P含量。     

The impact of shade on establishment of shrubs adapted to the high light irradiation of semi-arid environments

Light may be an important limiting resource that influences community structure of chenopod shrublands. As part of a larger study that aimed to determine the factors that influence chenopod community structure, the focus of this study was the influence of plant canopy on the growth and establishment of smaller plants. We therefore measured the height and cover of three chenopods (Enchylaena tomentosa, Maireana brevifolia and Maireana georgei) when growing within and outside of the canopy of Atriplex bunburyana under field conditions. All three chenopods had lower cover and E. tomentosa was taller when growing within the canopy of A. bunburyana in comparison to those growing outside of the canopy. The chenopods were then grown under three artificial shade regimes. Plant height, cover, biomass, relative leaf area and photosynthetic surface area measurements showed that each species responded differently to shade. E. tomentosa biomass was facilitated by shade. This was inferred by an increase in total plant biomass. M. brevifolia, in contrast, tolerated shade by increasing above-ground biomass allocation. M. georgei was adversely affected by the shade regimes: root biomass decreased in response to shade. Competition for light is, therefore, likely to influence chenopod community structure of semi-arid and arid environments.     

Responses of photosynthesis and water relations to rainfall in the desert shrub creosote bush (Larrea tridentata) as influenced by municipal biosolids

Responses of photosynthesis (P_n), stomatal conductance (g_s), pre-dawn leaf water potential (Ψ_lp) and leaf water content (ω_l) of creosote bush to 10 rainfall events in the Chihuahuan Desert were investigated. Infiltration of rainwater was manipulated by applying municipal biosolids. The responses of P_nand water relation parameters to rainfall (>10 mm) were mainly dependent upon drought severity: (1) following a moderate drought, P_n, g_s, Ψ_lpand ω_lrecovered to corresponding values of irrigated plants within 2 days after a 23-mm rainfall; (2) Ψ_lpand g_sresponded to a 15-mm rainfall within 2 days, following a 25-day drought, whereas responses of P_nand ω_lwere delayed for several days; (3) responses of P_n, g_s, Ψ_lpand ω_lto a 14·7-mm rainfall were all delayed for several weeks following a 110-day drought, but the delay was longer in P_n, g_sand ω_lthan in Ψ_lp. Creosote bush responded to small rainfall events (approximately 6 to 8 mm) with an increase in Ψ_lp, but without noticeable changes in g_sand P_n, suggesting a strong stomatal control of water loss even though xylem embolism was reduced. Biosolids applied at high rates (3·4 and 9 kgm⁻²) decreased the soil water by 2 to 4 mm following rainfall events, and this in turn delayed and decreased the responses of P_nand water relation parameters to rainfall.P_nand g_swere linearly related to ω_land exponentially related to Ψ_lp. With the generally coincidental responses of P_nor g_sand ω_lto rainfall, we concluded that the responses of P_nand g_sto rainfall were dependent on leaf rehydration which resulted from restored hydraulic conductance following drought.     

The influence of elevation, shrub species, and biological soil crust on fertile islands in the Mojave Desert, USA

We quantified soil nutrients and biological crust cover (bryophytes and lichens) under the canopies of three species of Mojave Desert shrubs and in interspaces between shrubs at three elevations to determine the effects of shrub species, soil crust, and elevation on islands of soil fertility. Means of pH, organic matter, total Kjeldahl nitrogen, nitrogen mineralization, and gravimetric soil moisture are significantly greater in soils under Ambrosia dumosa (Gray) Payne, Larrea tridentata Cov., and Coleogyne ramosissima Torr. than soils from adjacent interspace microhabitats. Although soil moisture and soil organic matter increase by a factor of 1.5 from the low elevation to the high elevation site, the ratio of shrub to interspace concentrations, or the difference in mean soil variables between shrubs and interspaces, is effectively constant and independent of elevation. Total bryophyte and lichen cover is relatively low (24.5%), however, there are 11 species of bryophytes and two species of lichens distributed across three elevations with the highest species richness and cover at the low-elevation site. Bryophyte and lichen cover is correlated with silt but is not related, consistently, to soil nutrients. Overall, the balance of processes controlling spatial aggregation of soil nutrients under shrubs is remarkably insensitive to potential differences in organic inputs among elevations, shrub species, and soil crust surfaces.     

Influence of shrubs on soil characteristics and their function in Sahelian agro-ecosystems in semi-arid Niger

Soil beneath shrubs form ‘fertile islands’ in fallow sites and millet fields in semi-arid Niger. To gain more information about this phenomena different shrub species in fallow sites following a gradient from 350–650 mm precipitation were examined. For each shrub two different areas were distinguished: an area under the canopy of the shrubs and an area in the nearby open land. Soil samples were taken from a depth of 0–10 cm and analysed for C_org, N_total, P_Bray, pH(H₂O), exchangeable cations, effective cation exchange capacity (ECEC) and soil texture. Significantly higher concentrations between 38–51% for C, N, P and 22% on ECEC for K⁺were found in the soil under the shrubs. The pH showed only slight but significant differences, whereas Al³⁺and H⁺rates on ECEC under the shrubs were increased by 44–55%. For Guiera senegalensis, the most common shrub of the studied area, enrichment ratios of most soil properties increased relatively more with increasing aridity. In general, enrichment ratios decreased with the age of the fallows, whereas concentrations showed no clear evolution. The chemical composition of the shrub litter seems to influence the degree of soil enrichment. The main step of fertile island formation takes place during the cultivation period by trapping wind-blown sediment. This work shows that shrubs are of vital importance for the accumulation of nutrients and maintenance of soil fertility within agro-ecosystems of Niger.     

Leaf-level physiological responses of Tamarix ramosissima to increasing salinity

Over the past century, the invasive halophytic shrub Tamarix ramosissima Ledeb. has increased in abundance and distribution in riparian ecosystems of western North America. These increases coincide with anthropogenic modification of river systems, which decrease the rate of periodic overbank flooding, leading to an increase in soil salinity. Increased soil salinity negatively impacts the physiology of native riparian tree species, but the impact of increased soil salinity on T. ramosissima physiology is incompletely known. To measure the impacts of soil salinity on T. ramosissima, we measured leaf-level responses across a broad range of surface-soil salinity concentrations at two sites in western Kansas. Photosynthesis at 2000 μmol m⁻² s⁻¹ (A₂₀₀₀), stomatal conductance to water (g_s), intercellular CO₂ concentration (C_i), and leaf δ¹³C showed little change over surface-soil salinities from 0.5 to 17.65 mmhos/cm. The small variation in leaf-level physiological responses suggests robust functioning of T. ramosissima across a broad range of surface-soil salinities. Leaf-level physiology and δ¹³C responses were assessed by canopy position, but responses were not significantly different. These results are among the first to show broad acclimation and robust physiological functioning for many leaf-level processes measured on mature trees grown across a wide surface-soil salinity gradient in the field.     

Water relations and hydraulic architecture of two Patagonian steppe shrubs: Effect of slope orientation and microclimate

On a local scale, topography influences microclimate, vegetation structure and the morpho-physiological attributes of plants. We studied the effects of microclimatic differences between NE- and SW-facing slopes on the water relations and hydraulic properties of two dominant shrubs of the Patagonian steppe in Argentina (Retanilla patagonica and Colliguaja integerrima). The NE-facing slope had higher irradiance and air saturation deficits and lower soil water availability and wind speed than the SW-facing slope. Predawn and midday Ψ_L and osmotic potentials were significantly lower in shrubs on the NE-facing slope. Osmotic adjustment and more elastic cell walls helped the plants to cope with a more xeric environment on NE-facing slope. Higher water deficits on NE-facing slope were partially compensated by a higher leaf and stem water storage. While stem hydraulic efficiency did not vary between slopes, leaf hydraulic conductance was between 40% and 300% higher on the NE-facing slope. Changes observed in leaf size and in SLA were consistent with responses to mechanical forces of wind (smaller and scleromorphic leaves on SW-facing slope). Morpho-physiological adjustments observed at a short spatial scale allow maintenance of midday Ψ_L above the turgor loss point and demonstrate that leaves are more responsive to microclimatic selective pressures than stems.     

Spatial and temporal variations in C3 and C4 plant abundance over the Chinese Loess Plateau since the last glacial maximum

Stable carbon isotopes of soil organic matter from 12 sites in the southern Chinese Loess Plateau are reviewed to examine spatial and temporal patterns of C₃ and C₄ plants in the arid to semiarid monsoonal region during three key periods - last glacial maximum, mid-Holocene, and modern. We have tentatively corrected the effects of atmospheric CO₂ concentrations and precipitation amounts on the δ¹³C endmembers for C₃ plants to reconstruct the relative proportion of C₄ plants because the δ¹³C values of C₃ plants are variable under different CO₂ and climate conditions. The results indicate that C₄ grasses increase from northwest to southeast spatially, which is consistent with present-day increasing precipitation and temperature patterns. This suggests that for a monsoon-dominant arid to semiarid region, such as the Loess Plateau, warm-season rainfall is a primary factor limiting C₄ plants growth, and thus C₄ grasses have been outcompeted by C₃ grasses/shrubs under cold-dry climate conditions. However, temporal fluctuations in atmospheric CO₂ concentration may also affect plant growth through altering water-use efficiency (WUE). Enhanced drought caused by decreased WUE due to low atmospheric CO₂ concentrations during glacial periods, combined with a cold-dry climate, leads to a decline in C₄ grasses, canceling out any advantages gained from lowered atmospheric CO₂ concentrations. To reconstruct accurately the abundance of C₃ and C₄ plants in an ecosystem and explore their controlling factors, process-based vegetation models integrating CO₂ and climatic parameters interactive with plant physiology are necessary.     

Light responses of Carex planostachys from various microsites in a Juniperus community

Juniperus communities are found on over 50 × 10⁶ ha in arid and semiarid habitats in southwestern North America. The drought tolerant sedge Carex planostachys occurs below the canopy in some of these communities. Cover and biomass of C. planostachys are high below the canopy and low in associated gaps. The purposes of this study were to investigate the temporal and spatial physiologic response of C. planostachys to abiotic changes, and determine it's light response characteristics from four contiguous microsites. Net photosynthesis was highest in spring when temperature was cooler and soil water higher, but low carbon uptake continued during summer drought. In addition, C. planostachys demonstrates a capacity to recover from extreme drought, despite water potential measured below ?9.0 MPa. Based on physiological light response curves and gas-exchange measurements, C. planostachys appears tolerant of shaded and full sun habitats. Light levels below the canopy were reduced compared to the gaps, but light saturation of C. planostachys did not change and net CO₂ uptake was only reduced slightly. Carbon uptake was coupled to light levels and not soil moisture. Observed differences in physiological attributes and variation in C. planostachys cover and biomass correspond to the presence or absence of the canopy. Low light compensation points, coupled with reduced respiratory demand, maximize photosynthetic gain in low light microsites. C. planostachys appears to acclimate across a range of light regimes, suggesting photosynthetic plasticity, allowing growth and survival in diverse light microhabitats. C. planostachys, tolerant of drought, appears anisohydric and demonstrates a capacity to acclimate to sun and shaded habitats, which could allow it to occur across a wider range of arid areas.     

The effect of radiation on seedling growth and physiology in four species of Prosopis L. (Mimosaceae)

Domestication of wild plants needs efforts focused particularly on the development of the theoretical basis of plant responses to environmental conditions. The objective of this study was to assess the effect of radiation on seedling growth and physiology of Prosopis alba, P. chilensis, P. flexuosa and P. glandulosa. Seedlings were grown in a greenhouse and randomly assigned to three light environments: full-sun, 52% sun, and 38% sun. No significant differences were found in the rate of leaf appearance and plant height 60 days after the light treatments began. Instantaneous CO₂uptake was not affected by the light environment during leaf development and did not differ among species. Variations in radiation availability resulted in significant differences in biomass accumulation, shoot/root ratio, protein and total non-structural carbohydrates contents. The practice of shading seedlings reduces the chances of survival ofProsopis due to an increase of the shoot/root ratio and a reduction of total biomass, stored carbohydrates, and the C:N balance.     

FINE SCALE VARIABILITY IN SOIL FROST DYNAMICS SURROUNDING CUSHIONS OF THE DOMINANT VASCULAR PLANT SPECIES (AZORELLA SELAGO) ON SUB‐ANTARCTIC MARION ISLAND

Through changing soil thermal regimes, soil moisture and affecting weathering and erosion processes plants can have an important effect on the physical properties and structure of soils. Such physical soil changes can in turn lead to biological facilitation, such as vegetation‐banked terrace formation or differential seedling establishment. We studied the fine scale variability in soil temperature and moisture parameters, specifically focusing on frost cycle characteristics around cushions of the dominant, vascular plant species, Azorella selago, on sub‐Antarctic Marion Island. The frost season was characterised by numerous low intensity and very shallow frost cycles. Soils on eastern cushion sides were found to have lower mean and maximum temperatures in winterthan soils on western cushion sides. In addition, lower variability in temperature was found on eastern cushion sides in winterthan on western cushion sides, probably as a result of higher wind speeds on western cushion sides and/or eastern, lee‐side snow accumulation. Despite the mild frost climate, extensive frost heave occurred in the study area, indicating that needle ice forms at temperatures above ?2°C. Our results demonstrate the effectiveness of frost pull as a heave mechanism under shallow frost conditions. The results highlight the importance of Azorella cushions in modifying site microclimates and of understanding the consequences of these modifications, such as potentially providing microhabitats. Such potential microhabitats are particularly important in light of current climate change trends on the island, as continued warming and drying will undoubtedly increase the need for thermal and moisture refugia.