Growth and reproduction of Antarctic vascular plants in response to warming and UV radiation reductions in the field

Along the west coast of the Antarctic Peninsula springtime ozone depletion events can lead to a two-fold increase in biologically effective UV-B radiation (UV-B_BE) and summer air temperatures have risen ≈1.5°C during the past 50 years. We manipulated levels of UV radiation and temperature around Colobanthus quitensis (a cushion-forming plant, Caryophyllaceae) and Deschampsia antarctica (a tussock grass) along the Peninsula near Palmer Station for two field seasons. Ambient levels of UV were manipulated by placing filters that either transmitted UV (filter control), absorbed UV-B (reducing diurnal levels of UV-B_BE by about 82%), or absorbed both UV-B and UV-A (reducing UV-B_BE and UV-A_BE by about 88 and 78%, respectively) on frames over naturally growing plants from November to March. Half the filters of each material completely surrounded the frames and raised diurnal and diel air temperatures around plants by an average of 2.3°C and 1.3°C, respectively. Reducing UV or warming had no effect on leaf concentrations of soluble UV-B absorbing compounds, UV-B absorbing surface waxes or chlorophylls. Warming had few effects on growth of either species over the first season. However, over the second field season warming improved growth of C. quitensis, leading to a 50% increase in leaf production (P < 0.10), a 26% increase in shoot production, and a 6% increase in foliar cover. In contrast, warming reduced growth of D. antarctica, leading to a 20% decline in leaf length, a 17% decline in leaf production (P < 0.10), and a 5% decline in foliar cover. Warming improved sexual reproduction in both species, primarily through faster development of reproductive structures and greater production of heavier seeds. Over the second field season, the percentage of reproductive structures that had reached the most developed (seed) stage in C. quitensis and D. antarctica was 20% and 15% higher, respectively, under warming. Capsules of C. quitensis produced 45% more seeds under warming and these seeds were 11% heavier. Growth of D. antarctica was improved when UV was reduced and these effects appeared to be cumulative over field seasons. Over the second season, tillers produced 55% more leaves and these leaves were 32% longer when UV-B was reduced. Tillers produced 137% more leaves that were 67% longer when both UV-B and UV-A were reduced. The effects of UV reduction were not as pronounced on C. quitensis, although over the second season cushions tended to be 17% larger and produce 21% more branches when UV-B was reduced, and tended to be 27% larger and produce 38% more branches when both UV-B and UV-A were reduced (P < 0.10). Few interactions were found between UV reduction and warming, although in the absence of warming, reducing UV led to slower development of reproductive structures in both species. The effects of warming and UV reduction were species specific and were often cumulative over the two field seasons, emphasizing the importance of long-term field manipulations in predicting the impacts of climate change. Received: 4 August 1998 / Accepted: 1 December 1998     

Current status of the Antarctic herb tundra formation in the Central Argentine Islands

Changes in the higher plant populations of the Argentine Islands over the last four to five decades have been central to developing an understanding of the likely biological responses to the globally exceptional rates of regional climate change, in particular warming, experienced along the western Antarctic Peninsula over the same period. In this study, we reassessed local populations and distribution of the two indigenous flowering plants on two islands in this archipelago, the grass Deschampsia antarctica and the pearlwort Colobanthus quitensis , in order to compare with previous partial and detailed surveys carried out by the British Antarctic Survey between 1963 and 1990. Our major finding was that the strong trend of recent increase in population size documented in 1990 has not continued, with current population sizes of both higher plants now being slightly lower than but still comparable with those recorded in the last survey in 1990. We discuss reasons underlying this, including possible limits imposed by the suitability of available habitat, and a recent plateauing of the local climate warming trend in comparison with that seen before the 1990 survey, with no significant short-term warming apparent in annual or seasonal meteorological data since 1990.     

Do Antarctic lichens modify microclimate and facilitate vascular plants in the maritime Antarctic? A comment to Molina‐Montenegro et al. ()

A recent article published by Molina‐Montenegro et al. (Journal of Vegetation Science24: 463) examines the association of Antarctic native plant and lichen species to the lichen Usnea antarctica on Fildes Peninsula, King George Island, maritime Antarctica. The authors report that on two sites, five out of 13 and four out of 11 species of lichens and mosses were spatially associated with U. antarctica, suggesting positive interactions between them. Although Deschampsia antarctica does not grow naturally associated with U. antarctica, Molina‐Montenegro et al. carried out a transplantation experiment to demonstrate that the macrolichen acts as a nurse plant, improving the survival of the grass. Serious conceptual and methodological discrepancies emerge from a critical evaluation of this study, challenging their conclusions. First, we suspect that the author confused some lichen taxa, and we also disagree with macrolichens being treated as cushion plants, because rootless, poikilohydric and poikilothermic thallophytes such as lichens are unable to create a stable, enclave‐like microhabitat as vascular cushion plants do. Indeed, a critical evaluation of some of the micro‐environmental parameters measured indicates that there are no biologically meaningful differences between the U. antarctica thalli and surrounding open areas. Second, the lack of consideration of the life history of the species under study leads to confusion when (a) referring to the succession sequence of species that colonize the studied area and (b) interpreting the putative distribution patterns promoted by Usnea versus the substrate preferences of associated species. Third, the authors intend to demonstrate experimentally that Usnea can facilitate the survival of D. antarctica plants, transplanting adult plants and not seedlings between the lichen thalli, and it is not clear how the grass was planted – between or within the lichens – as at both experimental sites the lichens grow on stones or rocks. Facilitative interactions are present in the Antarctic and may play a pivotal role in the structure and functioning of the fragile Antarctic ecosystems. However, more rigorous and well‐planned research is needed to assess its presence, importance and involved mechanisms.     

Effect of springtime solar ultraviolet-B radiation on growth of Colobanthus quitensis at Palmer Station, Antarctica

We examined the influence of solar ultraviolet‐B radiation (UV‐B; 280–315 nm) on the growth of Colobanthus quitensis plants by placing them under contrasting UV‐B filters at Palmer Station, along the Antarctic Peninsula. The filters reduced diurnal biologically effective UV‐B (UV‐B_BE) either by 83% (‘reduced UV‐B’) or by 12% (‘near‐ambient UV‐B’) over the 63 day experiment (7 November 1998–8 January 1999). Ozone column depletion averaged 17% during the experiment. Relative growth and net assimilation rates of plants exposed to near‐ambient UV‐B were 30 and 20% lower, respectively, than those of plants exposed to reduced UV‐B. The former plants produced 29% less total biomass, as a result of containing 54% less aboveground biomass. These reductions in aboveground biomass were mainly the result of a 45% reduction in shoot biomass, and a 31% reduction in reproductive biomass. Reductions in shoot biomass were owing to an 18% reduction in branch production by main shoots, while reductions in reproductive biomass were the result of a 19% reduction in individual capsule mass. Total plant leaf area was reduced by 19% under near‐ambient UV‐B, although total leaf biomass was unaffected because leaves had a greater specific leaf mass. The reduction in plant leaf area under near‐ambient UV‐B was attributable to: (1) production of 11% fewer leaves per main shoot system and plant, which resulted from an 18% reduction in branch production by main shoots. Leaf production per individual main shoot or branch was not affected; (2) shorter leaf longevity—main shoots contained 14% fewer green leaves at a given time; and (3) smaller individual leaves—leaf elongation rates were 14% slower and mature leaves were 13% shorter.     

Response of plants and the dominant microarthropod, Cryptopygus antarcticus, to warming and contrasting precipitation regimes in Antarctic tundra

We examined the influence of warming and supplemental precipitation on plant production and abundance of the dominant microarthropod, the springtail Cryptopygus antarcticus (Collembola), in tundra dominated by the vascular plants Colobanthus quitensis and Deschampsia antarctica along the Antarctic Peninsula. Tundra cores were placed in plots near Palmer Station where they were warmed with infrared heaters in combination with receiving supplemental precipitation. Diel canopy air and soil temperatures and air vapor pressure deficits in warmed plots were elevated 0.8 °C, 2.2 °C and 0.13 kPa, respectively. After two growing seasons, total aboveground plant production was greater under warming as a result of enhanced production by C. quitensis, which more than offset declines in moss biomass. Total aboveground plant production was also greater under supplemental precipitation primarily as a result of enhanced moss production. Total aboveground plant production was greatest under the combination of warming and supplemental precipitation, primarily as a result of enhanced C. quitensis production. C. antarcticus were more abundant in cores receiving supplemental precipitation and there was a strong treatment interaction; these springtails were most abundant in warmed cores receiving supplemental precipitation. Over 50% of the variability in the abundance of C. antarcticus could be explained by differences in aboveground plant biomass. However, plant production did not appear directly responsible for differences in C. antarcticus abundance; when we examined C. antarcticus abundance per unit of aboveground plant biomass, differences in its abundance among treatments were still apparent implying these differences were not the direct result of plant biomass. The responses of C. antarcticus were consistent with its known moisture and thermal preferences, suggesting that abiotic factors played a dominant role in controlling its abundance. Precipitation regime had large impacts on warming responses and these were species specific, illustrating the importance of future precipitation regimes in predicting system responses to warming.     

Warming increases aboveground plant biomass and C stocks in vascular-plant-dominated Antarctic tundra

We passively warmed tundra on the Antarctic Peninsula over four growing seasons and assessed its effect on dry mass and C and N stocks associated with the vascular plants Colobanthus quitensis (a cushion‐forming forb) and Deschampsia antarctica (a tussock grass), and mosses. Temperature treatments involved a warmed treatment that raised diurnal and diel canopy air temperatures by 2.3 and 1.3 °C, respectively, and a near‐ambient temperature treatment that raised diurnal and diel temperatures by 0.2 °C. These two different temperature regimes were achieved by wrapping filters around the frames to different extents and were nested within three UV treatments that filtered different solar UV wavebands. The experiment also included an ambient control treatment (unfiltered frames), and supplemental water and fertilizer treatments (applied to unfiltered frames). After four growing seasons, we collected cores of each vascular plant species and assessed the mass and C and N content of the aboveground current‐year biomass, the litter layer (which included nongreen live stems), and the organic soil horizon (which included roots). The thin nature of the organic soil horizon allowed us to sample this complete horizon and estimate near‐total ecosystem C and N stocks. A comparison of the warmed and near‐ambient temperature treatments found that warming led to greater aboveground biomass of C. quitensis, and more C in the aboveground biomass of both vascular plant species. Warming resulted in lower N concentrations of the aboveground biomass of both species. The water use efficiency of both species was greater under warming, based on their higher δ¹³C values. The mass of the litter layer under C. quitensis was greater under warming, and this layer contained more C and N and had a higher C : N ratio. The mass of the organic soil horizon under both species was greater under warming, and this horizon also contained more C and N. Warming also changed the species composition of the plant community – cover of C. quitensis increased while that of mosses declined. Warming resulted in the input of biomass into the system that had greater C : N ratios (and was likely more recalcitrant to decomposition) because (1) warming increased the C : N ratio of the biomass produced by both vascular plant species, (2) these inputs increased with warming because of greater biomass production, and (3) increases in C. quitensis cover led to greater biomass inputs by this species and its biomass had a greater C : N ratio than D. antarctica. Water or fertilizer supplements had few effects on aboveground biomass or C and N concentrations or pools, consistent with the relatively wet maritime climate and high soil nutrient levels of this system. Total C pools in the aboveground biomass, litter, and organic soil horizon were greater under warming. Warmed plots contained from 272 to 319 g m⁻² more C than plots under near‐ambient temperatures, corresponding to a 23–34% increase in ecosystem C.     

Vascular plants as bioindicators of regional warming in Antarctica

Monitoring selected populations of the only two native Antarctic vascular plant species (Colobanthus quitensis andDeschampsia antarctica) over a 27-year period has revealed a significant and relatively rapid increase in numbers of individuals and populations at two widely separated localities in the maritime Antarctic. There is strong evidence that this increase is a response to a warming trend in summer air temperatures, which has been evident throughout the region since the late 1940s, enhancing seed maturation, germination and seedling survival. This study provides the only known long-term monitoring data for any terrestrial organisms in Antarc-tica. Because their response to ameliorating conditions is more rapid than that of the dominant cryptogamic groups, Antarctic phanerogams may be useful bioindicators of climate change in West Antarctica.     

Photosynthesis of Marine Macroalgae from Antarctica: Light and Temperature Requirements

The photosynthetic performance of macroalgae isolated in Antarctica was studied in the laboratory. Species investigated were the brown algae Himantothallus grandifolius, Desmarestia anceps, Ascoseira mirabilis, the red algae Palmaria decipiens, Iridaea cordata, Gigartina skottsbergii, and the green algae Enteromorpha bulbosa, Acrosiphonia arcta, Ulothrix subflaccida and U. implexa. Unialgal cultures of the brown and red algae were maintained at 0°C, the green algae were cultivated at 10°C. I_K values were between 18 and 53 μmol m^?2 s^?1 characteristic or low light adapted algae. Only the two Ulothrix species showed higher I_K values between 70 and 74 μmol m^?2 s^?1. Photosynthesis compensated dark respiration at very low photon fluence rates between 1.6 and 10.6 μmol m^?2 s^?1. Values of α were high: between 0.4 and 1.1 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the brown and red algae and between 2.1 and 4.9 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the green algal species. At 0°C P_max values of the brown and red algae ranged from 6.8 to 19.1 μmol O₂ g^?1 FW h^?1 and were similarly high or higher than those of comparable Arctic-cold temperate species. Optimum temperatures for photosynthesis were 5 to 10°C in A. mirabilis, 10°C in H. grandifolius, 15°C in G. skottsbergii and 20°C or higher in D. anceps and I. cordata. P: R ratios strongly decreased in most brown and red algae with increasing temperatures due to different Q₁₀ values for photosynthesis (1.4 to 2.5) and dark respiration (2.5 to 4.1). These features indicate considerable physiological adaptation to the prevailing low light conditions and temperatures of Antarctic waters. In this respect the lower depth distribution limits and the northern distribution boundaries of these species partly depend on the physiological properties described here.     

Seasonal changes in temperature and nutrient control of photosynthesis, respiration and growth of natural phytoplankton communities

1. To investigate the influence of elevated temperatures and nutrients on photosynthesis, respiration and growth of natural phytoplankton assemblages, water was collected from a eutrophic lake in spring, summer, autumn, winter and the following spring and exposed to ambient temperature and ambient +2, +4 and +6 °C for 2 weeks with and without addition of extra inorganic nutrients. 2. Rates of photosynthesis, respiration and growth generally increased with temperature, but this effect was strongly enhanced by high nutrient availability, and therefore was most evident for nutrient amended cultures in seasons of low ambient nutrient availability. 3. Temperature stimulation of growth and metabolism was higher at low than high ambient temperature showing that long‐term temperature acclimation of the phytoplankton community before the experiments was of great importance for the measured rates. 4. Although we found distinct responses to relatively small temperature increases, the interaction between nutrient availability, time of the year and, thus, ambient temperature was responsible for most of the observed variability in phytoplankton growth, photosynthesis and respiration. 5. Although an increase in global temperature will influence production and degradation of organic material in lakes, the documented importance of ambient temperatures and nutrient conditions suggests that effects will be most pronounced during winter and early spring, while the remaining part of the growth season will be practically unaffected by increasing temperatures.     

ECOPHYSIOLOGICAL PERFORMANCE OF THE AEROTERRESTRIAL GREEN ALGA KLEBSORMIDIUM CRENULATUM (CHAROPHYCEAE,STREPTOPHYTA) ISOLATED FROM AN ALPINE SOIL CRUST WITH AN EMPHASIS ON DESICCATION STRESS1

Aeroterrestrial filamentous green algae of the genus Klebsormidium (Klebsormidiales, Streptophyta) are typical components of biological soil crusts, which occur worldwide in arid and semiarid habitats including alpine regions. In the present study, Klebsormidium crenulatum (Kütz.) Lokhorst was isolated from an alpine soil crust above the timberline of the Austrian Alps. Growth responses, photosynthetic performance, and desiccation tolerance were measured under controlled laboratory conditions. K. crenulatum exhibited optimal growth and the highest photosynthetic efficiency under relatively low photon fluence densities (30 and 21.9 μmol photons · m^?2 · s^?1, respectively), indicating low‐light requirements. It grew in a narrow range of salinities between 1.2 and 15 practical salinity units (psu), pointing to a pronounced stenohaline response pattern. Increasing temperatures from 5°C to 40°C led to different effects on photosynthetic oxygen evolution and respiratory oxygen consumption in K. crenulatum. While at low temperatures (5°C–10°C) photosynthesis was relatively high, respiration was not detectable or was at a very low level. Conversely, at the highest temperature of 40°C, photosynthesis was inhibited, and respiration unaffected, indicating strong differences in temperature sensitivity between both physiological processes. K. crenulatum was capable of photosynthesizing efficiently for up to 2.5 h under desiccation, followed by a decrease to 15% of the initial value after 3 h. Complete recovery took place within 2 h after rehydration. All ecophysiological data explain the widespread abundance of K. crenulatum in soil crusts of the alpine regions of the European Alps.     

Anatomical features and ultrastructure of Deschampsia antarctica (Poaceae) leaves from different growing habitats

BACKGROUND AND AIMS: The leaf anatomy and ultrastructure of Deschampsia antarctica (Poaceae) plants growing in three different habitats (a dry site in the Antarctic tundra, a wet site in a zone exposed to sea spray and a greenhouse) were investigated. The ultrastructure of the leaves of D. antarctica has not been studied before. METHODS: Semi-thin sections of the D. antarctica leaves were stained with toluidine blue and viewed using a light microscope. Ultra-thin sections stained with uranyl acetate and lead citrate were examined using a transmission electron microscope. KEY RESULTS: Plants growing in the Antarctic tundra and in a greenhouse had stronger xerophytic features than those growing at the seashore. The stress response of D. antarctica plants growing in the wet environment, exposed to high salinity and flooding, included: irregular mesophyll cells, large intercellular spaces in the parenchymatic layer, bulliform epidermal cells and vascular bundles surrounded with deformed outer and inner bundle sheaths of leaves. The highest number of sclerenchymatic fibres is characteristic of the leaves of plants growing in a greenhouse, whereas the smallest was of plants growing in a wet habitat. Stress conditions can disturb the formation of sclerenchymatic fibres. In plants growing in the Maritime Antarctic the chloroplasts of the mesophyll cells of leaves are of an irregular shape, with pockets or invaginations inside the organelles and outgrowths. Both of them make the surfaces of chloroplasts larger, and result in an increase in the amount of substances exchanged between the chloroplasts and cytoplasm or the other organelles. The leaf mesophyll cells of D. antarctica plants growing in Antarctica contain atypical structures including numerous vesicles of different sizes and concentrically arranged membranes. CONCLUSIONS: The anatomical and ultrastructural features of the leaf and their changes under stress conditions are considered in relation to the adaptations of D. antarctica to the climate conditions in the Maritime Antarctic.     

气候变暖对陆地植被-土壤生态系统的影响研究

由于自然因素及人类活动的长期影响,全球气候变化已经成为不容置疑的事实,并对陆地生态系统的植被及土壤产生了深远影响。陆地植被一土壤生态系统在全球气候变化中的反应与适应等过程已成为众多科学家所关注的问题。为更好地了解陆地植被一土壤生态系统对全球气候变化的响应机制,综述了气候变暖对植物的物候与生长、光合特征、生物量生产与分配,以及土壤呼吸等方面的影响,并对分析得到的结论进行了总结。分析指出,随着全球气候变暖,植物个体和群落特征以及土壤特性都会发生相应改变,高海拔地区的植被高度有增加趋势,而低海拔地区的植被可能出现矮化。然而,在以下方面还存有不确定性：（1）气候变暖导致的植被特征变化是否会减弱全球气候变化;（2）在较长时间尺度上气候变暖如何影响植物的物候和生长,特别是植物的体型;（3）高寒生态系统冬季土壤呼吸对气候变暖如何响应。     

增温和刈割对高寒草甸土壤呼吸及其组分的影响

评估土壤呼吸及其组分对增温等全球变化的响应对于预测陆地生态系统碳循环至关重要。利用红外线辐射加热器(Infrared heater)装置在青藏高原高寒草甸生态系统设置增温和刈割野外控制实验。通过测定2018年生长季(5—9月)土壤呼吸和异养呼吸,探究增温和刈割对土壤呼吸及其组分的影响。研究结果表明:(1)单独增温使土壤呼吸显著增加31.65%(P<0.05),异养呼吸显著增加27.12%(P<0.05),土壤自养呼吸没有显著改变(P>0.05);单独刈割对土壤呼吸和自养呼吸没有显著影响(P>0.05),单独刈割刺激异养呼吸增加32.54%(P<0.05);(2)增温和刈割之间的交互作用对土壤呼吸和异养呼吸没有显著影响(P>0.05),但是对自养呼吸的影响是显著的(P<0.05),土壤呼吸和异养呼吸的季节效应显著(P<0.05);(3)土壤呼吸及其组分与土壤温度均成显著指数关系,与土壤湿度呈显著的正相关关系(P<0.05),处理影响它们的响应敏感性。本研究表明青藏高原东缘高寒草甸土壤碳排放与气候变暖存在正反馈。