Habitat-specific responses in the flowering phenology and seed set of alpine plants to climate variation: implications for global-change impacts

The timing of the snowmelt is a crucial factor in determining the phenological schedule of alpine plants. A long-term monitoring of snowmelt regimes in a Japanese alpine area revealed that the onset of the snowmelt season has been accelerated during the last 17 years in early snowmelt sites but that such a trend has not been detected in late snowmelt sites. This indicates that the global warming effect on the snowmelt pattern may be site-specific. The flowering phenology of fellfield plants in an exposed wind-blown habitat was consistent between an unusually warm year (1998) and a normal year (2001). In contrast, the flowering occurrence of snowbed plants varied greatly between the years depending on the snowmelt time. There was a large number of flowering species in the fellfield community from mid- to late to late June and from mid- to late July. The flowering peak of an early-melt snowbed plant community was in the middle of the flowering season and that of a late-melt snowbed community was in the early flowering season. These habitat-specific phenological patterns were consistent between 1998 and 2001. The effects of the variation in flowering timing on seed-set success were evaluated for an entomophilous snowbed herb, Peucedanum multivittatum, along the snowmelt gradient during a 5-year period. When flowering occurred prior to early August, mean temperature during the flowering season positively influenced the seed set. When flowering occurred later than early August, however, the plants enjoyed high seed-set success irrespective of temperature conditions if frost damage was absent. These observations are probably explained based on the availability of pollinators, which depends not only on ambient temperature but also on seasonal progress. These results suggest that the effects of climate change on biological interaction may vary depending on the specific habitat in the alpine ecosystem in which diverse snowmelt patterns create complicated seasonality for plants within a very localized area.     

Assimilation and translocation of nitrogen and carbon in Curcuma alismatifolia Gagnep

Curcuma or Siam tulip (Curcuma alismatifolia Gagnep.) is an ornamental flowering plant with two underground storage organs, rhizomes and storage roots. Characteristics of N and C assimilation and transport in curcuma were investigated. The plants were treated with ¹⁵NH₄⁺ + ¹⁵NO₃^? and ¹³CO₂ at 10, 13 or 21 weeks after planting. Plants were sampled at several stages up to 32 weeks. The C stored in old storage roots was used rapidly during the first 10 weeks; after which N stored in old rhizomes and old storage roots were used. The daily gain in C depending on photosynthesis was remarkably high between 10 and 21 weeks. However, the daily gain in N was relatively constant throughout the growth period. The ¹⁵N absorbed at 10 weeks was initially accumulated in leaves and roots, but some was transported to flowering organs at 13 weeks. At harvest, 41% of ¹⁵N was recovered in new rhizomes and 17% in new storage roots. After ¹³CO₂ exposure at 10 and 13 weeks, the distribution of ¹³C among organs was relatively constant in subsequent stages. When given ¹³CO₂ at 21 weeks, a large amount of labelled C was recovered in new storage roots and new rhizomes at harvest. Both new rhizomes and new storage roots stored N and C, however, rhizomes played a more important role in supplying N, while storage roots provided C.     

Carbohydrate storage and use in an alpine population of the perennial herb, Oxytropis sericea

I tested hypotheses for ecological roles of storage carbohydrates in perennating organs (roots and branches) of alpine Oxytropis sericea, a leguminous herb. In naturally growing plants, total nonstructural carbohydrates achieved their maximal concentration in the fall, declined during winter, and reached minimal levels immediately after growth initiation in the spring. Experimental manipulation of carbon sink-source relations through shading of leaves of reproductive plants revealed that the normally unused portion of these carbohydrates is largely available for withdrawal. In another experiment, plants subjected to carbohydrate depletion through shading suffered decreased leaf growth after winter dormancy and had a lower probability of flowering and decreased inflorescence biomass. The dependence of reproductive growth on stored carbohydrates, however, was limited to its initial stages, because accumulation of storage carbohydrates occurred simultaneously with inflorescence expansion, flowering, and fruiting. Moreover, the whole-plant photosynthetic rate, estimated from gas exchange measurements also peaked at the time of inflorescence growth. To address whether stored reserves allow compensatory regrowth following defoliation, plants were subjected to experimental removal of leaves and inflorescences. Defoliated O. sericea partly regrew the lost leaves but withdrawal of stored carbohydrates was limited. Similarly, in a second defoliation experiment where infructescences were left intact, the plants used little stored carbohydrate and only partly compensated for fruit growth. However, carbohydrate accumulation was negatively affected by defoliation. While the ecological importance of stored nonstructural carbohydrates cannot be attributed to any function in isolation, winter respiration, leaf regrowth after winter, and early reproductive growth in O. sericea all depend to a significant extent on stored reserves. Maintaining a large storage pool may protect these functions in years when carbon status is less favorable than during this study. Received: 13 May 1998 / Accepted: 24 November 1998     

Carbohydrate storage in meadow plants and its depletion after disturbance: do roots and stem-derived organs differ in their roles?

Storage of carbohydrates in organs protected from disturbance is an important adaptation of plants in disturbed habitats. We carried out a field experiment involving 31 herbaceous plant species in two cultural meadows to find out whether roots or belowground stem-derived organs (stem bases, stem tubers and rhizomes) are the main storage organs, to study how reserves accumulate in individual organs in the long term (growing season) and to ascertain whether meadow abandonment affects the distribution of carbohydrate reserves in plants. We also conducted a 22-day pot experiment with four meadow plant species to determine how removal of roots and aboveground parts affects the use of carbohydrates stored in roots and stem-derived organs in the short term. From the long-term perspective of the field experiment, mowing had a positive effect on the concentration of carbohydrate reserves. From the short-term perspective of the pot experiment, however, the effect on concentration and pools of carbohydrates was negative. In the field experiment, carbohydrate concentrations before winter were generally higher than in mid-season, and more often higher in roots than in stem-derived organs. Roots and stem-derived organs of plants in the pot experiment were depleted similarly after both types of disturbance. Our results indicate a need for including both types of belowground plant organs in future studies of the carbon economy of plants from disturbed habitats.     

Fine-scale dynamics of rhizomes in a grassland community

Spatial dynamics in grassland communities are constrained by the belowground spatial distribution of roots and rhizomes. Their dynamics are difficult to measure as underground data collection tends to be destructive and cannot be repeated at the same plot over time. We investigated rhizome dynamics indirectly by examining rhizome spatial structure on long‐term grassland study plots where aboveground shoot counts have been recorded using a fine‐scale grid over nine years. Number of rhizome apical ends, basal ends and total rhizome length of both live and dead rhizomes were obtained from the data by scanning rhizomes and processing them by GIS vectorization. These rhizome variables were correlated with the above‐ground shoot counts in grid cells over varying temporal lags. There was a general decrease in the intensity of correlation between live rhizomes and shoot counts with increasing time lags. Correlation of dead rhizomes increased with increasing time lag, reaching a maximum after several years, and then declined. Species differed strongly in the change of rhizome‐shoot counts correlation over varying time lags. These differences were used to infer rhizome growth dynamics, namely rhizome growth rate and lifespan, and rhizome mean decomposition time. The species involved differed in all these traits. Mean rhizome growth rate ranged from 0.2 (Polygonum bistorta) to 3.3 cm ur^?1 (Deschampsia flexuosa); mean rhizome lifespan ranged from 5 yr (Anthoxanthum alpinum) to over 8 yr (Nardus stricta) and mean decomposition time from one growing season (Anthoxanthum) to 7 yr (Polygonum). Presence of dead rhizomes below living rhizomes or aboveground shoots was taken as an indication of fine‐scale replacements between species. These were highly non‐random, with some species pairs replacing significantly more frequently. These differences in rhizome growth parameters underlie different strategies of horizontal growth and dieback between species. These can serve as one of the mechanisms of species replacements and contribute to the fine‐scale coexistence of species.     

The Utilization of Carbohydrate and Nitrogen Reserves by Taxus During Its Spring Growth Period

The spring growth and the utilization of carbohydrate and nitrogen reserves in this growth was studied in Taxus media cv. Hicksii plants 0, 2, 4 and 6 weeks after the plants started growing in the spring. The effect of nitrogen applied the previous season on the storage and utilization of the carbohydrate and nitrogen reserves during spring growth was determined. The plants were separated into buds (all new growth), stems, needles (those produced the previous season) and roots and analyzed for changes in total nitrogen, basic and non-basic amino acids, total available carbohydrate, sugars, hemicelluloses, organic acids and chlorophyll. The bulk of the soluble nitrogen reserves were stored as arginine in the stems and old needles. With the onset of spring growth, arginine nitrogen was converted to other amino acids which accumulated in the new growth (buds). The roots, stems and needles of plants grown under high nitrogen levels always contained more total nitrogen than those grown under low nitrogen levels. The bulk of the carbohydrate reserves were stored as hemicelluloses. The plants grown under high nitrogen levels utilized the bulk of the carbohydrate reserves from the roots and smaller amounts from the stems and old needles, while plants grown under low nitrogen levels used only the reserves in the roots. In the low nitrogen plants, carbohydrates accumulated in the needles and stems. Both the carbohydrate and nitrogen reserves were important in the dry weight increase due to spring growth. However, the nitrogen reserves were the limiting factor and the high nitrogen plants grew twice as much, produced more chlorophyll, and utilized more nitrogen and carbohydrate reserve in spring growth than low nitrogen plants. The additional chlorophyll allowed the production of more carbohydrates and these additional carbohydrates were used in increased growth rates, while in the low nitrogen plants the carbohydrate produced was less and accumulated within the plant.     

Adaptive significance of nitrogen storage in Bistorta bistortoides,an alpine herb

Summary Studies were conducted to examine the importance of nitrogen storage to seasonal aboveground growth in the alpine herb Bistorta bistortoides. Stored reserves accounted for 60% of the total nitrogen allocated to the shoot during the growing season. The stored nitrogen was equally partitioned between preformed buds of the shoot and the roots/rhizome. Reliance on stored N was similar in populations of a 105-day growing season site and of a 75-day growing season site. Contrary to our initial hypothesis, stored nitrogen reserves were not used to extend the growing season of this species into the late-spring when soils are still cold, and saturated with snow-melt water. The time at which stored nitrogen was used to initiate shoot growth coincided with the time of root initiation, rapid soil warming, and near maximum soil concentrations of NO _inf3 ^sup– and NH _inf4 ^sup+ . Thus, nitrogen demand and soil nitrogen supply were both high at the same time. The importance of nitrogen storage in this species appeared to be in satisfying the high demand of simultaneous vegetative and reproductive growth during the early-growing season after soils thawed. The initiation of rapid leaf and inflorescence growth occurred in mid-June in both sites. The maximum pool size of shoot nitrogen (maximum nitrogen demand) occurred only 12 days later in the long season site, and 28 days later in the short season site. The early-season utilization of nitrogen stores allows plants of this species to initiate reproductive allocation at the same time vegetative tissues are exhibiting maximal growth rates. By releasing vegetative and reproductive growth from competition for nitrogen, seeds could mature early in the alpine growing season, before the frost probability sharply increases in mid-August.     

Relationship between flowering time and fruit set of the entomophilous alpine shrub,Rhododendron aureum (Ericaceae), inhabiting snow patches

Flowering phenology, fruit set, and pollinator frequency of the ericaceous alpine dwarf shrub, Rhododendron aureum, were compared among three quadrats arranged along a snowmelt gradient on a slope. Bagging and self-pollination experiments showed that R. aureum was physiologically self-compatible, but pollinator visitation highly enhanced fruit set. Depending on the lateness of snowmelt, flowering time varied from mid-June to late July among quadrats. Pollinator visitation increased as the flowering season progressed, and fruit set was significantly higher in the quadrat with latest snowmelt. Emasculation experiments suggested that later flowering might promote cross-pollination. Thus, later flowering was advantageous for effective pollination. On the other hand, later-flowering plants often failed to set fruits because of the onset of autumn frost and snow before fruit maturation.     

Changes in carbohydrate contents of <Emphasis Type="Italic">Zantedeschia</Emphasis> leaves under gibberellin-stimulated flowering

The effect of gibberellic acid (GA₃) on the carbohydrate accumulation in relation to vegetative growth of Zantedeschia ‘Black Magic’ plants undergoing transition to flowering was investigated. In response to GA treatment the carbohydrate level increased independently of earlier stimulation of shoot emergence. Under vegetative growth stage the content of reducing sugars of leaf blades was 2.5-fold higher than in control plants, and suggests the stimulation of photosynthetic activity. The changes observed during the flowering, in principle noted in petiole tissues, support the GA-effect on assimilate transport to the sink organs. Moreover, the high level of non-structural carbohydrates in petiole tissues, in particular reducing sugars, can be an effect of photosynthetic activity of these organs and/or essential for osmoregulation and high turgor pressure. The results indicate that apart from the influence on the shoot emergence, the GAs may stimulate the photosynthetic activity from the beginning of shoot growth and are thus responsible for the enhancement of callas flower yield.     

The importance of nitrogen and carbohydrate storage for plant growth of the alpine herb Veratrum album

We examined whether nitrogen (N) and carbohydrates reserves allow Veratrum album, an alpine forb, to start spring growth earlier than the neighbouring vegetation and to survive unpredictable disturbances resulting in loss of above-ground biomass. * Seasonal dynamics of plant reserves, soil N availability and vegetation growth were monitored. Veratrum album shoots were experimentally removed when carbohydrate reserves were at a seasonal minimum and the subsequent changes in biomass and reserves were compared with those in control plants. Reserves did not give V. album a competitive advantage in spring; however, they did function as a buffer against the impact of calamities. Shoot removal resulted in significantly lower root dry weight, higher N concentration in rhizome and roots and lower starch concentrations in rhizome and roots but no plant mortality was observed. Veratrum album used stored N reserves to supplement N uptake and establish high leaf N concentrations, which facilitated a rapid refilling of depleted carbohydrate reserves. The primary function of N reserves appears to be to allow V. album to complete the growing cycle in as short a period as possible, thus minimizing exposure to above-ground risks.     

Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrate reserves

We examined interspecific and intraspecific variation in tree seedling survival as a function of allocation to carbohydrate reserves and structural root biomass. We predicted that allocation to carbohydrate reserves would vary as a function of the phenology of shoot growth, because of a hypothesized tradeoff between aboveground growth and carbohydrate storage. Intraspecific variation in levels of carbohydrate reserves was induced through experimental defoliation of naturally occurring, 2-year-old seedlings of four northeastern tree species –Acer rubrum, A. saccharum, Quercus rubra, and Prunus serotina– with shoot growth strategies that ranged from highly determinate to indeterminate. Allocation to root structural biomass varied among species and as a function of light, but did not respond to the defoliation treatments. Allocation to carbohydrate reserves varied among species, and the two species with the most determinate shoot growth patterns had the highest total mass of carbohydrate reserves, but not the highest concentrations. Both the total mass and concentrations of carbohydrate reserves were significantly reduced by defoliation. Seedling survival during the year following the defoliation treatments did not vary among species, but did vary dramatically in response to defoliation. In general, there was an approximately linear relationship between carbohydrate reserves and subsequent survival, but no clear relationship between allocation to root structural biomass and subsequent survival. Because of the disproportionate amounts of reserves stored in roots, we would have erroneously concluded that allocation to roots was significantly and positively related to seedling survival if we had failed to distinguish between reserves and structural biomass in roots. Received: 14 December 1999 / Accepted: 2 June 1999     

Seasonal changes in carbohydrate reserves in mature northern<Emphasis Type="Italic"> Populus tremuloides</Emphasis> clones

To assess the changes in seasonal carbohydrate status of Populus tremuloides, sugar and starch concentrations were monitored in roots, stem xylem and phloem and branches of ten different clones. Time of root growth was assessed by extraction of roots from in-growth cores collected five times during the season. Overall the results showed that the main period of root growth in these northern clones was shifted from spring to late summer and fall likely due to the microclimatic conditions of the soil. This increase in root growth was associated with a decline in total non-structural carbohydrate content in the roots during this period. This study also found that the carbohydrate reserves in these clones were being stored as close as possible to the organs of annual growth (leaves and roots). At the time of leaf flush, the largest reduction in stored carbohydrates (3% of dry weight) was observed in the branches of the trees, compared to a slight decline in the stem and roots. Starch and sugar reserves in most tissues were very low in early summer. This suggests that reserves that might be used for the regrowth of foliage after insect defoliation or other disturbances, are relatively small compared to the portion that is needed for maintenance and typical growth developments such as leaf flush.     

Mycorrhizal C costs and nutritional benefits in developing grapevines

Arbuscular mycorrhizal (AM) C-costs in grapevines were investigated. Dormant vines rely on stored C for initial growth. Therefore AM colonisation costs would compete with plant growth for available C reserves. One-year-old grapevines, colonised with Glomus etunicatum (Becker and Gerdemann), were cultivated under glasshouse conditions. The C-economy and P utilisation of the symbiosis were sequentially analysed. AM colonisation, during the 0–67 day growth period, used more stem C relative to root C, which resulted in lower shoot growth. The decline in AM colonisation during the period of 67–119 days coincided with stem C replenishment and higher shoot growth. Construction costs of AM plants and root C allocation increased with root P uptake. The efficiency of P utilisation was lower in AM roots. The reliance of AM colonisation on stem C declined with a decrease in colonisation, providing more C for the refilling of stem carbohydrate reserves and shoot growth. Once established, the AM symbiosis increased P uptake at the expense of refilling of root C reserves. Although higher root C allocation increased plant construction costs, AM roots were more efficient at P utilisation.     

Vulnerability of phenological synchrony between plants and pollinators in an alpine ecosystem

The relationship between flowering phenology and abundance of bumble bees (Bombus spp.) was investigated using 2 years of phenological data collected in an alpine region of northern Japan. Abundance of Bombus species was observed along a fixed transect throughout the flowering season. The number of flowering species was closely related to the floral resources for pollinators at the community scale. In the year with typical weather, the first flowering peak corresponded to the emergence time of queen bees from hibernation, while the second flowering peak corresponded to the active period of worker bees. In the year with an unusually warm spring, however, phenological synchrony between plants and bees was disrupted. Estimated emergence of queen bees was 10 days earlier than the first flowering date owing to earlier soil thawing and warming. However, subsequent worker emergence was delayed, indicating slower colony development. The flowering season finished 2 weeks earlier in the warm-spring year in response to earlier snowmelt. A common resident species in the alpine environment, B. hypocrita sapporoensis, flexibly responded to the yearly fluctuation of flowering. In contrast, population dynamics of other Bombus species were out of synchrony with the flowering: their frequencies were highest at the end of the flowering season in the warm-spring year. Therefore, phenological mismatch between flowers and pollinators is evident during warm years, which may become more prevalent in a warmer climate. To understand the mechanism of phenological mismatch in the pollination system of the alpine ecosystem, ground temperature, snowmelt regime, and life cycle of pollinators are key factors.     

The role of roots and cotyledons as storage organs in early stages of establishment in Quercus crispula: a quantitative analysis of the nonstructural carbohydrate in cotyledons and roots

Quercus seedlings have hypogeal cotyledons and tap roots, both of which act as storage organs. The importance of the storage function in the two organs may change as the seedling develops. Therefore, changes in carbohydrate reserves in cotyledons and roots of Q. crispula grown under 40 % and 3 % of full light from shoot emergence to the completion of the first leaf flush were monitored. In addition, a shoot-clipping treatment was performed to examine the relative contribution of the cotyledons and tap roots to resprouting. Cotyledons maintained large amounts of nonstructural carbohydrates during shoot development, and carbohydrates were still present in the cotyledons during the final phase of leaf flush. In addition, a notable increase in the amount of carbohydrates was observed in tap roots before leaf flush at both light levels. Since root development occurred before leaf flush, even in plants grown under 3 % light, the carbohydrate found in them presumably originated from seed reserves and was translocated to roots as storage reserves. When shoots were clipped at the leaf flushing stage, the amount of carbohydrate decreased only in the cotyledons after resprouting, suggesting that cotyledons act as the main storage organs during shoot development stages. However, it could be advantageous as a 'risk avoidance strategy' for the seedlings to store reserves in both cotyledons and roots, since cotyledons may be removed by predators during shoot development.     

Timing of reproductive and vegetative development in Saxifraga oppositifolia in an alpine and a subnival climate

Morphogenesis of floral structures, dynamics of reproductive development from floral initiation until fruit maturation, and leaf turnover in vegetative short-stem shoots of Saxifraga oppositifolia were studied in three consecutive years at an alpine site (2300 m) and at an early- and late-thawing subnival site (2650 m) in the Austrian Alps. Marked differences in the timing and progression of reproductive and vegetative development occurred: individuals of the alpine population required a four-month growing season to complete reproductive development and initiate new flower buds, whereas later thawing individuals from the subnival sites attained the same structural and functional state within only two and a half months. Reproductive and vegetative development were not strictly correlated because timing of flowering, seed development, and shoot growth depended mainly on the date of snowmelt, whereas the initiation of flower primordia was evidently controlled by photoperiod. Floral induction occurred during June and July, from which a critical day length for primary floral induction of about 15 h could be inferred. Preformed flower buds overwinter in a pre-meiotic state and meiosis starts immediately after snowmelt in spring. Vegetative short-stem shoots performed a full leaf turnover within a growing season: 16 (+/-0.8 SE) new leaves per shoot developed in alpine and early-thawing subnival individuals and 12 (+/-1.2 SE) leaves in late-thawing subnival individuals. New leaf primordia emerged continuously from snowmelt until late autumn, even when plants were temporarily covered with snow. Differences in the developmental dynamics between the alpine and subnival population were independent of site temperatures, and are probably the result of ecotypic adaptation to differences in growing season length.     

Seasonal changes in pollen limitation and femaleness along the snowmelt gradient in a distylous alpine herb,Primula modesta

Flowering phenology of alpine plants is strongly determined by the timing of snowmelt, and the conditions of pollination of widely distributed plants vary greatly during their flowering season. We examined the reproductive success of the distylous alpine herb, Primula modesta, along the snowmelt gradient under natural conditions, and compared it with the result of artificial pollination experiments. In addition, the compositions and visit frequencies of pollinators to the flower of P. modesta were examined during the flowering period. The pin and thrum plants of P. modesta growing at the same site have an equal ability to produce seeds if a sufficient amount of legitimate pollen grains are deposited on the stigma surface. However, under natural conditions, their seed‐set success was often (even if not always) restricted by pollen limitation, and the functional gender of the pin and thrum plants biased to the female and male, respectively, associated with their growing sites. These variations were not ascribed to resource limitation nor biased morph ratio but to the seasonal changes in pollination situations, a replacement of pollinator types from long‐ to short‐tongued pollinators resulted in unidirectional pollen transfer from long stamens (thrum plants) to long styles (pin plants). The functional gender specialization may enhance the evolution of dioecy from heterostyly, but the severe pollen limitation may cause the breakdown of heterostyly into homostyly. To consider the evolutionary pathway of heterostylous plants, an accumulation of the empirical data is required demonstrating how phenological synchrony between plants and pollinators is decided and to what degree this relationship is stable over years, along with estimates of selection and gene flow in individual plants.     

Seasonal changes in shoot regrowth potential in Calamagrostis canadensis

Summary A series of laboratory experiments was conducted to examine seasonal change in shoot regrowth potential following disturbance in Calamagrostis canadensis. On several dates during the 1988 and 1989 growing seasons, soil cores were collected from field sites dominated by this grass. Shoot regrowth from cores after clipping at the soil surface was monitored under dark or light laboratory conditions at 20°C. seasonal changes in field concentrations of total nonstructural carbohydrate and nitrogen in rhizomes largely accounted for the observed seasonal change in etiolated regrowth potential of shoots in laboratory experiments. In contrast, shoot regrowth potential in the light showed a very different seasonal pattern. The ratio of shoot biomass regrowth 20 d after clipping in the light versus dark treatment showed a gradual seasonal decrease from 12:1 in the early May experiment to near 1:1 in the September experiment. However, the rate of photosynthesis of regrowing shoots in the light was highest in experiments conducted late in the growing season. This may indicate a strong seasonal decrease in the proportion of current photosynthate of regrowing shoots that is allocated to new shoot growth. Alternatively, mobilization of rhizome carbohydrate reserves for shoot regrowth may have been inhibited during the re-establishment of photosynthesis in the light treatment. Either mechanism would explain why shoot regrowth in the light is poorly correlated with levels of belowground carbohydrate reserves, even under controlled laboratory conditions.     

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.     

天祝高寒珠芽蓼草甸初级生产力的研究I.生物量动态及光能转化率

甘肃天祝高寒珠芽蓼草甸5月20日左右返青。地上生物量的变化呈单峰曲线,最大值在8月22日,干物质为548.39g／m2(489.06g／m2去灰分物质;下同);净第一性生产力为481.05g／m2·a干物质。地下生物量很大,6一9月平均接近6kg／m2,呈单谷曲线变化,最低值出现在7月20日,为4556.87g／m2干物质。地上部分最大生长率出现在月平均气温只有8—10℃的返青后一个月,平均绝对生长率为5.89g／m2·d干物质,平均相对生长率为0.152g／g·d干物质。春季地上部分的最大生长率与活根的很大消耗联系在一起。地上部分对太阳总辐射的转化率为0.155%,对生理辐射的转化率为0.316%,对≥0℃-≤0℃生长期的生理辐射的转化率为0.692%。地上部分在生长的第一个月对总辐射的表观转化率最高,平均为0.57%。