高山植物的抗寒抗冻特性

综述了高山植物光合作用对冰冻低温胁迫的反应和适应特性,阐述了高山植物呼吸作用的低温适应特点;对高山植物抗冻机制作了详细分析和讨论。     

高山植物光合机构耐受胁迫的适应机制

植物的光合作用是易受环境影响的重要生理过程之一.高山植物作为生长在特定极端环境(低温/强辐射)下的植物群体,其光合器官在形态结构和生理功能上形成了抵御强辐射和低温胁迫的特殊适应机制.但由于较高的生境异质性,高山植物的光保护适应机制存在较大的差异.光保护适应机制与光合作用密切关联,影响植物的碳同化能力和生物量的形成能力.本文对近年来国内外有关高山植物光合器官叶绿体的形态、超微解剖结构及光合机构光保护适应机理的研究进展进行了综述,并提出了今后高山植物光合作用生理适应性研究的方向.     

高山植物的形态、结构和生理特征

高山植物是在高海拔地区或树线以上高山气候条件下生长的植物物种的总称,是植物长期适应高山恶劣环境而高度特化的结果。高山植物的类群主要包括多年生禾草、莎草、非禾草的垫状植物、苔藓、地衣等。有些高山植物也是药用植物。高山植物为了生存和繁殖种群,必须适应严酷的高山环境,包括低温、干旱、强紫外辐射和较短的生长季。从植株形态、器官解剖结构、光合作用、元素利用等方面阐述高山植物的基本特征及其适应高山环境的内在机制。在全球气候变化的大环境下,研究高山植物对环境的响应和适应性具有重要的理论意义。     

高山植物的光合生理特性研究进展

高山植物的光合作用受强辐射、低温和干旱环境的影响。近年来,大气CO_2浓度上升和全球气候变暖的趋势日益明显,影响着高山植物的光合生理。本文综述了强辐射、低温和干旱等高山环境因子以及全球气候变化引起的大气CO_2和温度上升对高山植物光合特性的影响,并提出未来高山植物光合生理的研究热点主要是开展不同地域的高山植物光合特性研究,环境因子交互作用对高山植物光合特性的影响研究,不同植物的光合特性对全球气候变化响应的差异,模拟土壤有效养分含量增高对高山植物光合特性的影响,建立数学模型预测全球气候变化对高山植物动态的影响以及通过长期定位研究探索不同生长阶段高山树木的光合特性。     

青藏高原高山植物的形态和解剖结构及其对环境的适应性研究进展

高山植物是一类生长于树线以上至雪线的山地植物。揭示高山植物适应环境的形态和结构特征及其内在机制,对研究全球气候变化下,植物对环境的响应和适应具有重要的理论意义。然而,国内在高山植物功能生态学的研究上相对薄弱,已有研究主要集中在对青藏高原高山植物的报道上。结合国外高山植物的相关研究报道,从形态和解剖结构两个方面对青藏高原高山植物的研究进展进行了综述,重点阐述了高山植物的形态、解剖结构及其与环境的适应性关系。植株矮小（有的呈垫状）、叶片小而厚、具有通气组织、栅栏组织多层、机械组织发达、虫媒花性状、线粒体数量多和叶绿体基粒片层少等是这一地区高山植物普遍具有的形态和结构特征。高山植物形成上述结构的特异性是高山特殊综合生态环境长期作用的结果。同时,也是高山植物对高山环境的高度适应。最后,对这一领域存在的问题以及未来研究的重点和方向进行了探讨。目的是引起国内研究者的关注,促进我国高山植物功能生态学的研究与发展。     

低温锻炼对绵头雪莲花组织培养苗抗寒性及抗氧化酶活性的影响

高山植物绵头雪莲花(Saussurea laniceps HandMazz.)组织培养苗经2℃低温锻炼后抗寒性明显增强,SOD、CAT、POD活性明显提高,可溶性蛋白质、脯氨酸含量升高,而可溶性糖含量没有明显变化.脱锻炼期间可溶性蛋白质含量仍高于低温锻炼期间的水平,脱锻炼后抗氧化酶活性略有上升或保持稳定状态.这些变化是绵头雪莲花组织培养苗适应低温逆境的生理生化基础.     

几种高山植物叶绿体淀粉粒的变化特征

利用透射电镜对生长于青藏高原东北部达坂山（海拔3900m）的5种高山植物叶绿体超微结构进行了观察。结果发现,在所研究的5种高山植物叶绿体中,淀粉粒数量均较多,淀粉粒呈长椭圆形或圆形,沿叶绿体长轴分布。在珠芽蓼的叶绿体中,淀粉粒的电子密度内外不均匀,外周电子密度低,中央电子密度高。在其余4种高山植物中,淀粉粒的电子密度均较低。另外,在这5种高山植物叶绿体中还出现了脂质小球。其类囊体均出现了不同程度的膨大现象。研究表明,高山植物叶绿体中淀粉粒的这种变化是对逆境的一种适应,是青藏高原特殊生态条件长期胁迫的结果。     

高山植物繁殖策略的研究进展

高山地区通常被认为是陆地上最为极端的生境之一,但却拥有许多形态特化的植物和较高的物种多样性。高山植物如何在严酷的环境中实现成功繁殖,这一问题倍受研究者们的关注。本文综合了国内外高山植物在资源分配、花形态对非生物环境因子的响应、动物传粉及其适应机制、果实和种子及克隆繁殖等繁殖策略方面的文献。为应对低温多雨雪的恶劣环境,一些高山植物采取花向日性、花冠闭合及花序保温结构等繁殖策略。高山植物的传粉者类群也发生了改变,主要为蜂类和蝇类。熊蜂(Bombusspp.)传粉的高效性,减少了高山环境对植物传粉造成的不利影响。当传粉者不可得时,植物不仅通过延迟自交和自助自交等机制来提供繁殖保障,还借助克隆繁殖及其他传粉机制(风媒或风虫媒)来维持种群的繁衍。依赖动物传粉的高山植物,可以采取增加繁殖构件的资源分配、加大"广告"投入以及较大的花展示或较长的花寿命来提高传粉者的拜访几率,以及借助泛化的花结构和选择合适的开花时间等策略来提高繁殖成功率。此外,大部分高山植物产生干果且具有持久的种子库,有利于种子的传播以及种子寻找萌发及幼苗生长的最佳外界环境。在今后的研究中,可着重探讨以下几个问题:(1)非生物环境因子对花形态的选择;(2)季节变化与繁殖策略;(3)群落水平上植物与传粉者的关系;(4)高山生态系统对全球变暖的响应。     

达坂山蚤缀和裸茎金腰叶绿体超微结构的研究

对高山植物达坂山蚤缀和裸茎金腰叶绿体超微结构的研究表明,叶绿体和线粒体镶嵌很紧密;基粒和基质类囊体膨大,膨大的类囊体呈梭形或圆形,没有发现脂质小球的存在;基粒叠垛程度很低,平均每个基粒类囊体的片层数为5．4。这2种高山植物叶绿体超微结构上的特征是青藏高原特殊的生态条件,包括低温、低气压、强辐射影响的结果。     

基于数字标本探究绿绒蒿属植物花期的变化

近年来,随着全球气温的明显升高,气候变化引起的全球变暖受到人们的广泛关注.气候变暖会使植物物候发生改变,花期的变化能够直观地反映出植物对周围环境的适应,尤其是高山植物对气候变化更为敏感,因此探究高山植物的花期更具有代表性.为了解植物在气候变暖背景下花期的变化特征,并制定出合理的植物保护措施,该文以典型的高山植物绿绒蒿属...     

Drought increases the freezing resistance of high-elevation plants of the Central Chilean Andes

Freezing temperatures and summer droughts shape plant life in Mediterranean high-elevation habitats. Thus, the impacts of climate change on plant survival for these species could be quite different to those from mesic mountains. We exposed 12 alpine species to experimental irrigation and warming in the Central Chilean Andes to assess whether irrigation decreases freezing resistance, irrigation influences freezing resistance when plants are exposed to warming, and to assess the relative importance of irrigation and temperature in controlling plant freezing resistance. Freezing resistance was determined as the freezing temperature that produced 50 % photoinactivation [lethal temperature (LT₅₀)] and the freezing point (FP). In seven out of 12 high-Andean species, LT₅₀ of drought-exposed plants was on average 3.5 K lower than that of irrigated plants. In contrast, most species did not show differences in FP. Warming changed the effect of irrigation on LT₅₀. Depending on species, warming was found to have (1) no effect, (2) to increase, or (3) to decrease the irrigation effect on LT₅₀. However, the effect size of irrigation on LT₅₀ was greater than that of warming for almost all species. The effect of irrigation on FP was slightly changed by warming and was sometimes in disagreement with LT₅₀ responses. Our data show that drought increases the freezing resistance of high-Andean plant species as a general plant response. Although freezing resistance increases depended on species-specific traits, our results show that warmer and moister growing seasons due to climate change will seriously threaten plant survival and persistence of these and other alpine species in dry mountains.     

THE ECOLOGY OF ARCTIC AND ALPINE PLANTS

‘How are plants adapted to the low temperatures and other stresses of arctic and alpine environments ?’ At present it is not possible to answer this question completely. Much work remains to be done, particularly on low-temperature metabolism, frost resistance, and the environmental cues and requirements for flowering, dormancy, regrowth, and germination. However, in brief, we can say that plants are adapted to these severe environments by employing combinations of the following general characteristics: 1. Life form: perennial herb, prostrate shrub, or lichen. Perennial herbs have greatest part of biomass underground. 2. Seed dormancy: generally controlled by environment; seeds can remain dormant for long periods of time at low temperatures since they require temperatures well above freezing for germination. 3. Seedling establishment: rare and very slow; it is often several years before a seedling is safely established. 4. Chlorophyll content: in both alpine and arctic ecosystems not greatly different on a land-area basis from that in temperate herbaceous communities. Within a single species there is more chlorophyll in leaves of arctic populations than in those of alpine populations. 5. Photosynthesis and respiration: (a) These are at high rates for only a few weeks when temperatures and light are favourable. (b) Optimum photosynthesis rates are at lower temperatures than for ordinary plants; rates are both genetically and environmentally controlled with phenotypic plasticity very marked. (c) Dark respiration is higher at all temperatures than for ordinary plants; rate is both genetically and environmentally controlled, with phenotypic plasticity very pronounced, i.e. low-temperature environment increases the rate at all temperatures. (d) Alpine plants have higher light-saturation values in photosynthesis than do arctic or lowland plants; light saturation closely tied to temperature. (e) There is some evidence that alpine plants can carry on photosynthesis at lower carbon dioxide concentrations than can other plants. (f) Annual productivity is low, but daily productivity during growing season can be as high as that of most temperate herbaceous vegetation. Productivity can be increased by temperature, nutrients, or water. 6. Drought resistance: most drought stress in winter in exposed sites is due to frozen soils and dry winds. It is met by decreased water potentials, higher concentrations of soluble carbohydrates, and closed stomates. Little drought resistance in snowbank plants. Alpine plants adapted to summer drought stress can carry on photosynthesis at low water potentials; alpine or arctic plants of moist sites cannot do this. 7. Breaking of dormancy: controlled by mean temperatures near or above 0° C., and in some cases by photoperiod also. 8. Growth: very rapid even at low positive temperatures. Respiration greatly exceeds photosynthesis in early re-growth of perennials. Internal photosynthesis may occur in hollow stems of larger plants during early growth. Nitrogen and phosphorus often limiting in cold soil. 9. Food storage: characteristic of all alpine and arctic plants except annuals. Carbohydrates mostly stored underground in herbaceous perennials. Lipids in old leaves and stems of prostrate evergreen shrubs. Depleted in early growth, and usually restored after flowering. 10. Winter survival: survival and frost resistance are excellent after hardening. Cold resistance closely tied to content of soluble carbohydrates, particularly raffinose. 11. Flowering: flower buds are pre-formed the year before. Complete development and anthesis dependent upon temperature of the flowering year and also, in some cases, upon photoperiod. 12. Pollination: mostly insect-pollinated in alpine regions and even in Arctic, but to a lesser extent. Wind-pollination increasingly more important with increasing latitude. Diptera more important than bees in the Arctic and in the highest mountains. 13. Seed production: opportunistic, and dependent upon temperature during flowering period and latter half of growing season. 14. Vegetative reproduction: by rhizomes, bulbils, or layering. More common and important in Arctic than in alpine areas. 15. Onset of dormancy: triggered by photoperiod, low temperatures, and drought. Dormant plant extremely resistant to low temperatures.     

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO₂ levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO₂ enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO₂ and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO₂ levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.     

Adaptations to low temperature in high altitude insects from Mount Kenya

Abstract. 1. The strategies for low temperature survival in insects on Mount Kenya were investigated. The insects were collected from their natural habitats and their supercooling points and low temperature tolerances determined.
2. Most insects showed no special adaptations to low temperature survival and seem to depend on spending the cold nights in protected habitats, such as beneath stones and fallen trunks of plants, as well as within the wet frills of dead leaves of alpine plants, where they are protected by the heat released from freezing water.
3. Some insects, e.g. Collembola, aphids and a curculionid beetle, which live in relatively unprotected habitats, had low supercooling points, allowing them to remain unfrozen when exposed to low night temperatures. A nucleator free diet is apparently essential for the survival of such species.
4. Two species of curculionid beetles were found to withstand freezing down to -7C. These beetles had nucleating agents in their haemolymph and higher supercooling points than most of the other species studied.
5. A moderate freezing tolerance was found in larvae of a midge that lives in the watery liquid between the leaves of Senecio brassica .     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.     

Foliar freezing resistance of Australian alpine plants over the growing season

We assessed the freezing resistance of leaves ex situ of 25 Australian alpine plant species. We compared the freezing resistance of forb, graminoid and shrub species from three alpine summits of different altitudes; from a low altitude site just above treeline, to a fully alpine tundra site. Foliar freezing resistance (LT₅₀) in spring varied from ?5.9°C to ?18.7°C and standardized LT₅₀ values within species were significantly related to site altitude. Additionally, when comparing all the species in the study, freezing resistance was significantly related to site; the LT₅₀ of samples from a low‐altitude summit (1696 m) were significantly lower than those of samples from mid‐ (1805 m) and high‐altitude (1860 m) summits. The LT₅₀ of juvenile foliage did not differ significantly from that of adult foliage. Shrubs were highly resistant to freezing. At the highest summit, we examined the course of seasonal freezing resistance from early summer to early autumn across three alpine plant communities that differed in the time of natural snowmelt; from sheltered (snowpatch) to exposed (open heath). No differences in freezing resistance over the growing season were detected for exposed or sheltered communities and there were no consistent trends indicating frost hardening over the growing season. Overall, the common Australian alpine species we investigated appear well adapted to freezing conditions throughout the snow‐free growing season. We have no evidence to suggest that freezing temperatures soon after snowmelt in spring are especially damaging to the alpine plants at these summits.     

Ecological and evolutionary responses of an arctic plant to variation in microclimate and soil

The arctic and alpine regions are predicted to experience some of the highest rates of climate change, and the arctic vegetation is expected to be especially sensitive to such changes. Understanding the ecological and evolutionary responses of arctic plant species to changes in climate is therefore a key objective. Geothermal areas, where natural temperature gradients occur over small spatial scales, and without many of the confounding environmental factors present in latitudinal and other gradient studies, provide a natural experimental setting in which to examine the response of arctic–alpine plants to increasing temperatures. To test the ecological and evolutionary response of the circumpolar alpine bistort Persicaria vivipara to temperature, we collected plant material and soil from areas with low, intermediate and high soil temperatures and grew them at three different temperatures in a three-factorial growth chamber experiment. At higher experimental soil temperatures, sprouting was earlier and plants had more leaves. Sprouting was earlier in soil originating from intermediate temperature and plants had more leaves when grown in soil originating from low temperatures. We did not find evidence of local adaptation or genetic variation in reaction norms among plants originating from areas with low, intermediate and high soil temperature. Our findings suggest that the alpine bistort has a strong plastic response to warming, but that differences in soil temperature have not resulted in genetic differentiation. The lack of an observed evolutionary response may, for example, be due to the absence of temperature-mediated selection on P. vivipara, the low rate of sexual recombination, or high levels of gene flow balancing differences in selection. When placed within the context of other studies, we conclude that arctic–alpine plant species often show strong plastic responses to spring warming, while evidence of evolutionary responses varies among species.     

Microsite characteristics of Muhlenbergia richardsonis (Trin.) Rydb., an alpine C4 grass from the White Mountains, California

C₄ plants are uncommon in cold environments and do not generally occur in the alpine tundra. In the White Mountains of California, however, the C₄ grass Muhlenbergia richardsonis is common in the alpine zone at 3,300-3,800 m, with the highest population observed at 3,960 m (13,000 feet) above sea level. This is the highest reported C₄ species in North America and is near the world altitude limit for C₄ plants (4,000-4,500 m). Above 3,800 m, M. richardsonis is largely restricted to southern slope aspects, with greatest frequency on southeast-facing slopes. In open tundra, M. richardsonis formed prostrate mats with a mean height of 2.5 cm. Neighboring C₃ grasses were two to three times taller. Because of its short stature, leaf temperature of M. richardsonis was greatly influenced by the boundary layer of the ground, rising over 20°C above air temperature in full sun and still air and over 10°C above air temperature in full sun and wind velocity of 1-4 m s^-1. Thus, although air temperatures did not exceed 15°C, midday leaf temperatures of M. richardsonis were routinely between 25°C and 35°C, a range favorable to C₄ photosynthesis. At night, leaf temperature of M. richardsonis was often 5-12°C below air temperature, resulting in regular exposure to subzero temperatures and frosting of the leaves. No visible injury was associated with exposure to freezing night temperatures. The presence of M. richardsonis in the alpine zone demonstrates that C₄ plants can tolerate extreme cold during the growing season. The localization to microsites where leaf temperatures can exceed 25°C during the day, however, indicates that even when cold tolerant, C₄ plants still require periods of high leaf temperature to remain competitive with C₃ species. In this regard, the prostrate growth form of M. richardsonis compensates for the alpine climate by allowing sufficient heating of the leaf canopy during the day.     

Thermal tolerance of tropical and temperate alpine plants suggests that ‘mountain passes are not higher in the tropics’

Aim

Tolerance of species to extreme temperatures largely determines their distribution and vulnerability to climate change. We examined thermal tolerance in tropical and temperate alpine plants, testing the hypotheses that: (a) temperate plants are resistant to more extreme temperatures and have an overall wider thermal tolerance breadth (TTB); (b) TTB in temperate plants is wider than TTB in tropical plants during the entire growing season; (c) resistance to frost and heat varies during the season in temperate plants but not in tropical plants; (d) TTB of a species predicts its latitudinal range.

Location

Tropical (Ecuador, Bolivia) and temperate (USA, Austria) mountains.

Time period

Four periods of the growing season (2014, 2016–2019).

Major taxa

Ninety-six vascular plant species.

Methods

We employed the electrolyte leakage method to estimate the temperature resistance, that is, the temperature at which 50% tissue injury (Lt50) occurs in leaves. We used phylogenetic linear mixed-effect models in a Bayesian framework to test for differences between the plant groups.

Results

Temperate and tropical plants do not differ in their temperature resistance. The four hypotheses are rejected since: (a) temperate plants do not have significantly wider overall TTB compared to tropical plants, (b) TTB of temperate plants is wider than TTB of tropical plants only at the end of the temperate summer, (c) seasonal acclimation is observed in both plant groups, (d) the latitudinal range of the plants is not related to TTB.

Main conclusions

The lack of TTB differences between temperate and tropical alpine plants is consistent with trends observed in ectothermic animals, which suggests a general latitudinal pattern in high-elevation poikilotherm organisms. Limited acclimation capacity to cope with long freezing exposures restricts the occurrence of tropical alpine species to thermally aseasonal environments making them particularly vulnerable to climate change.