Contrasting growth changes in two dominant species of a Mediterranean shrubland submitted to experimental drought and warming

BACKGROUND AND AIMS: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. METHODS: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. KEY RESULTS: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6 degrees C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. CONCLUSIONS: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term.     

Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7 years (1999–2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area.     

Experimental drought and warming decrease diversity and slow down post-fire succession in a Mediterranean shrubland

Plant community recovery (species richness, diversity and composition) of a post-fire Mediterranean shrubland was monitored over a seven year period (1998–2005) under experimental drought and warming that simulated the environmental conditions forecast for this area in the coming decades. Species richness and Shannon's index were positively correlated with accumulated precipitation in the growing season and both variables were negatively affected by reduced water availability in drought plots. The relative abundance of the different species in both treatments was linearly correlated with their relative abundance in control plots. Moreover, we found species-specific responses to treatments. Drought and warming treatment reduced the competitive ability of the obligate seeder tree Pinus halepensis against native resprouter shrubs and consequently, the transformation from shrub to pine tree dominated vegetation was slowed down. Conversely, the water use strategy of Globularia alypum may allow this species to maintain a dominant position in drought plots. Therefore, future drier and warmer conditions in Mediterranean areas may severely affect plant community recovery after a disturbance, due to the existence of both abundance-dependent and species-specific responses that may change inter-specific competitive relationships.     

Precipitation-dependent flowering of Globularia alypum and Erica multiflora in Mediterranean shrubland under experimental drought and warming, and its inter-annual variability

Background and Aims

Relationships between autumn flowering, precipitation and temperature of plant species of Mediterranean coastal shrublands have been described, but not analysed experimentally. These relationships were analysed for two species of co-occurring, dominant, autumn-flowering shrubs, Globularia alypum and Erica multiflora, over 4 years and in experimentally generated drought and warming conditions. The aim was to improve predictions about the responses and adaptations of flowering of Mediterranean vegetation to climate change.

Methods

Beginning of anthesis and date of maximum flowering intensity (‘peak date’) were monitored over 4 years (2001–2004) on a garrigue land type in the noth-east of the Iberian Peninsula. Two experimental treatments were applied, increased temperature (+0·73°C) and reduced soil moisture (–17%) relative to untreated plots.

Key Results

Flowering of Globularia alypum and Erica multiflora differed greatly between years depending on the precipitation of the previous months and the date of the last substantial rainfall (>10 mm). Globularia alypum flowered once or twice (unimodal or bimodal) as the result of differences in the distribution and magnitude of precipitation in late-spring and summer (when floral buds develop). The drought treatment delayed and decreased flowering of Globularia alypum in 2001 and delayed flowering in 2002. Warming extended the period between the beginning of flowering and the end of the second peak for autumn flowering in 2001 and also increased peak intensity in 2002. Flowering of Erica multiflora was unaffected by either treatment.

Conclusions

Autumn flowering of Globularia alypum and Erica multiflora is more dependent on water availability than on temperature. Considerable inter-annual plasticity in the beginning of anthesis and peak date and on unimodal or bimodal flowering constitutes a ‘safe strategy’ for both species in relation to varying precipitation and temperature. However, severe changes in precipitation in spring and summer may severely affect flowering of Globularia alypum but not Erica multiflora, thus affecting development/structure of the ecosystem if such conditions persist.Key words: Globularia alypum, Erica multiflora, autumn flowering, drought, global warming, Mediterranean     

Drought and warming induced changes in P and K concentration and accumulation in plant biomass and soil in a Mediterranean shrubland

A field experiment involving drought and warming manipulation was conducted over a 6-year period in a Mediterranean shrubland to simulate the climate conditions projected by IPCC models for the coming decades (20% decreased soil moisture and 1°C warming). We investigated P and K concentration and accumulation in the leaves and stems of the dominant species, and in soil. Drought decreased P concentration in Globularia alypum leaves (21%) and in Erica multiflora stems (30%) and decreased K concentration in the leaves of both species (20% and 29%, respectively). The general decrease of P and K concentration in drought plots was due to the reduction of soil water content, soil and root phosphatase activity and photosynthetic capacity that decreased plant uptake capacity. Warming increased P concentration in Erica multiflora leaves (42%), but decreased it in the stems and leaf litter of Erica multiflora and the leaf litter (33%) of Globularia alypum, thereby demonstrating that warming improved the P retranslocation and allocation from stem to leaves. These results correlate with the increase in photosynthetic capacity and growth of these two dominant shrub species in warming plots. Drought and warming had no significant effects on biomass P accumulation in the period 1999–2005, but drought increased K accumulation in aboveground biomass (10 kg ha⁻¹) in Globularia alypum due to the increase in K concentration in stems. The stoichiometric changes produced by the different responses of the nutrients led to changes in the P/K concentration ratio in Erica multiflora leaves, stems and litter, and in Globularia alypum stems and litter. This may have implications for the nutritional value of these plant species and plant–herbivore relationships. The effects of climate change on P and K concentrations and contents in Mediterranean ecosystems will differ depending on whether the main component of change is drought or warming.     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Changes in the onset of spring growth in shrubland species in response to experimental warming along a north–south gradient in Europe

Aim To test whether the onset of spring growth in European shrublands is advanced in response to the warmer conditions projected for the next two decades by climate models, and, if there is a change, whether it differs across Europe. Location The studied sites spanned a broad north–south European gradient with average annual temperatures (8.2–15.6 °C) and precipitation (511–1427 mm). Methods ‘Bud break’ was monitored in eight shrub and grass species in six European sites under control and experimentally warmer conditions generated by automatic roofs covering vegetation during the night. Results Species responsive to increased temperatures were Vaccinium myrtillus and Empetrum nigrum in Wales, Deschampsia flexuosa in Denmark, Calluna vulgaris in Netherlands, Populus alba in Hungary and Erica multiflora in Spain. Although the acceleration of spring growth was the commonest response to warming treatments, the responses at each site were species specific and year dependent. Under experimental warming 25% of cases exhibited a significantly earlier onset of the growing season and 10% had a significantly delayed onset of vegetative growth. No geographical gradient was detected in the experimental warming effects. However, there was a trend towards a greater dominance of phenological advances with more intense the warming treatments. Above 0.8 °C warming, only advancements were recorded. Main conclusions Our results show that warmer temperatures projected for the next decades have substantial potential effects on the phenology of the spring growth of dominant species in different European shrublands, with a dominant trend towards advancements the more intense the warming is. However, our study also demonstrates the overall difficulties of applying simple predictive relationships to extrapolate the effects of global change on phenology. Various combinations of environmental factors occur concurrently at different European sites and the interactions between different drivers (e.g. water and chilling) can alter phenology significantly.     

Tissue-water relations of two co-occurring evergreen Mediterranean species in response to seasonal and experimental drought conditions

Tissue-water relations were used to characterize the responses of two Mediterranean co-occurring woody species (Quercus ilex L. and Phillyrea latifolia L.) to seasonal and experimental drought conditions. Soil water availability was reduced 15% by partially excluding rain throughfall and lateral flow (water runoff). Seasonal and experimental drought elicited physiological and morphological adaptations other than osmotic adjustment: both species showed large increases in cell-wall elasticity and decreased saturated-to-dry-mass ratio. Increased elasticity (lower elastic modulus) resulted in concurrent decreases in relative water content at turgor loss. In addition, P. latifolia showed significant increases in apoplastic water fraction. Decreased saturated-to-dry-mass ratio and increased apoplastic water fraction were accompanied by an increased range of turgor maintenance, which indicates that leaf sclerophyllous traits might be advantageous in drier scenarios. In contrast, the degree of sclerophylly (as assessed by the leaf mass-to-area ratio) was not related to tissue elasticity. An 15% reduction in soil water availability resulted in significant reductions in diameter growth when compared to control plants in both species. Moreover, although P. latifolia underwent larger changes in tissue water-related traits than Q. ilex in response to decreasing water availability, growth was more sensitive to water stress in P. latifolia than in Q. ilex. Differences in diameter growth between species might be partially linked to the effects of cell-wall elasticity and turgor pressure on growth, since Q. ilex showed higher tissue elasticity and higher intrinsic tolerance to water deficit (as indicated by lower relative water content at turgor loss) than P. latifolia.     

Resilience of Mediterranean shrubland to a severe drought episode: the role of seed bank and seedling emergence

Extreme climate events, such as severe drought episodes, may induce changes in vegetation if they induce species‐specific adult mortality and changes in the seedling recruitment pattern. In 2005 a severe drought occurred in Doñana National Park (south Spain) causing extensive shrubland mortality. Over the following years we monitored the soil seed bank and seedling emergence via a gradient of canopy dieback induced by the drought episode. The canopy dieback corresponded to an increase in emergence of seedlings of woody species in 2007, probably because of the reduced competition induced by canopy loss. The soil seed bank of woody species sampled in 2008 was less abundant on plots with a higher proportion of dead vegetation, probably because of depletion of the seed bank as a result of the increased germination in the previous year and also as a result of a reduction in seed supply in these sites. Accordingly, in 2009 we detected reduced emergence of woody species on plots that had suffered the greatest shrub mortality. We failed to find any significant changes in patterns of the soil seed bank and seedling emergence of short‐lived herbaceous species, indicating greater resilience in these types of species. This study highlights the resilience of Mediterranean shrublands to climate fluctuations at one extreme of the variability characteristic of these ecosystems. An increase in the frequency of severe drought episodes – increasingly probable under the new climate conditions – does have the potential, however, to induce changes in vegetation, especially in woody communities that need more time to replenish their seed banks.     

Effects of seasonal and decadal warming on soil enzymatic activity in a P‐deficient Mediterranean shrubland

Soil enzymes are central in the response of terrestrial ecosystems to climate change, and their study can be crucial for the models’ implementation. We investigated for 1 year the effects of warming and seasonality on the potential activities of five soil extracellular enzymes and their relationships with soil moisture, phosphorus (P) concentration, and other soil parameters in a P‐limited Mediterranean semiarid shrubland. The site was continuously subjected to warming since 1999, and we compared data from this study to analogous data from 2004. Warming uniformly increased all enzymes activities, but only when a sufficient amount of soil water was available. Seasonality unevenly altered enzyme activities, thus affecting enzymatic stoichiometry. P deficiency affected enzymatic stoichiometry, favoring the activities of the phosphatases. The effect of warming was stronger in 2014 than 2004, excluding the hypothesis of acclimation of rhizospheric responses to higher temperatures and suggesting that further increases in extracellular enzymatic activities are to be expected if sufficient water is available. Climatic warming will likely generally stimulate soil enzymatic activities and accelerate nutrient mineralization and similar ecological processes such as the production and degradation of biomass and changes in community composition, but which will be limited by water availability, especially in Mediterranean soils in summer. Winters in such ecosystems will benefit from a general increase in activity and production, but biological activity could even decrease in summer, potentially leading to a negative overall balance of nutrient mineralization. This study suggests that a general increase in activity due to warming could lead to faster mineralization of soil organic matter and water consumption in colder climates, until one of these factors in turn becomes limiting. Such trade‐offs between water and temperature in relation with enzyme activity should be considered in biogeochemical models.     

Effects of long-term experimental night-time warming and drought on photosynthesis, <Emphasis Type="Italic">F</Emphasis>v/<Emphasis Type="Italic">F</Emphasis>m and stomatal conductance in the dominant species of a Mediterranean shrubland

We conducted a night-time warming and drought field experiment for 7 years (1999–2005) in a Mediterranean shrubland. We focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree Pinus halepensis L. and the final years to study the effects of the experimental night-time warming and drought on Fv/Fm, photosynthesis, and stomatal conductance. Warming treatment increased mean air temperature and mean soil temperature through the years by an average of 0.7 and 0.9°C respectively, and drought treatment reduced soil moisture through the years by an average of 19%. Warming tended to increase photosynthetic rates in E. multiflora, G. alypum and P. halepensis mostly in the cold seasons, when plants were more limited by temperature, as shown by the lowest values of Fv/Fm being detected in winter in the three studied species. A negative effect of warming was only detected for E. multiflora in summer 2003. Drought treatment generated different responses of net photosynthetic rates depending on the species, season and year. Stomatal conductance showed the same pattern as photosynthesis for the three studied species, displaying seasonal and inter-annual variability, although with an overall negative effect of drought for P. halepensis. Photosynthetic rates decreased significantly in the dry winter 2005 and spring 2005 in comparison to the same seasons of 2003 and 2004. There were positive correlations between the photosynthetic rates in different seasons for E. multiflora, G. alypum and P. halepensis and the soil moisture of the week prior to measurements. The great variation in the photosynthetic rates was thus explained in a significant part by soil moisture levels. The lowest Fv/Fm values usually corresponded with lowest stomatal conductances suggesting that drought stress could be associated to stress by low temperatures in winter.     

Impact of drought and increasing temperatures on soil CO2 emissions in a Mediterranean shrubland (gariga)

In arid and semiarid shrubland ecosystems of the Mediterranean basin, soil moisture is a key factor controlling biogeochemical cycles and the release of CO₂ via soil respiration. This is influenced by increasing temperatures. We manipulated the microclimate in a Mediterranean shrubland to increase the soil and air night-time temperatures and to reduce water input from precipitation. The objective was to analyze the extent to which higher temperatures and a drier climate influence soil CO₂ emissions in the short term and on an annual basis. The microclimate was manipulated in field plots (about 25 m²) by covering the vegetation during the night (Warming treatment) and during rain events (Drought treatment). Soil CO₂ effluxes were monitored in the treatments and compared to a control over a 3-year period. Along with soil respiration measurements, we recorded soil temperature at 5 cm depth by a soil temperature probe. The seasonal pattern of soil CO₂ efflux was characterized by higher rates during the wet vegetative season and lower rates during the dry non-vegetative season (summer). The Warming treatment did not change SR fluxes at any sampling date. The Drought treatment decreased soil CO₂ emissions on only three of 10 occasions during 2004. The variation of soil respiration with temperature and soil water content did not differ significantly among the treatments, but was affected by the season. The annual CO₂ emissions were not significantly affected by the treatments. In the semi-arid Mediterranean shrubland, an increase of soil CO₂ efflux in response to a moderate increase of daily minimum temperature is unlikely, whereas less precipitation can strongly affect the soil processes mainly limited by water availability.     

Seasonal patterns of terpene content and emission from seven Mediterranean woody species in field conditions

The seasonal pattern of terpene content and emission by seven Mediterranean woody species was studied under field conditions. Emission rates were normalized at 30°C and 1000 μmol·m·s PFD (photosynthetic photon flux density). Bupleurum fruticosum, Pinus halepensis, and Cistus albidus stored large amounts of terpenes (0.01-1.77% [dry matter]) with maximum values in autumn and minimum values in spring. They emitted large amounts of terpenes (2-40 μg·g DM·h), but with no clear seasonal trend except for Cistus albidus, which had maximum values in spring and minimum values in autumn. The nonstoring species Arbutus unedo, Erica arborea, Quercus coccifera and Quercus ilex also emitted large amounts of terpenes (0-40 μg·g DM·h) and also tended to present maximum emission rates in spring, although this trend was significant only for A. unedo. At the seasonal scale, emission rates did not follow changes in photosynthetic rates; instead, they mostly followed changes in temperature. From autumn to spring, the least volatile monoterpenes such as limonene were emitted at highest rates, whereas the most volatile monoterpenes such as α-pinene and β-pinene were the most emitted in summer. The monoterpene emission rates represented a greater percentage of the photosynthetic carbon fixation in summer (from 0.51% in Arbutus unedo to 5.64% in Quercus coccifera) than in the rest of the seasons. All these seasonality trends must be considered when inventorying and modeling annual emission rates in Mediterranean ecosystems.     

两种圆柏属植物不同季节显微和超微结构变化与耐寒性的关系

以祁连圆柏和圆柏为材料,研究了圆柏属植物的抗寒性与解剖结构变化,以及淀粉与可溶性糖粉转化的关系.结果表明,祁连圆柏和圆柏叶片的上下表皮细胞外覆盖一层角质膜,叶肉细胞间隙大,形成发达的通气组织.在生长季节,二者叶绿体内均积累了很多淀粉粒,在冷冻休眠季节淀粉粒消失或变小,同时叶片可溶性糖分含量增加,祁连圆柏的可溶性糖分增幅高于圆柏;而休眠季节圆柏叶绿体受到低温伤害,脂质球数目增多,出现脂质小滴,部分叶绿体变形,祁连圆柏类囊体的结构受到影响较小,脂质球数量少,叶绿体形状没有发生明显的变化.因此,祁连圆柏和圆柏抗寒性与发达的通气组织和淀粉粒生长季节的积累有关,在低温胁迫下祁连圆柏叶绿体表现出高的稳定性,通过可溶性糖分的调节力强,耐寒性增强.     

Variation and response to experimental warming in a New Zealand cushion plant species

Cold adapted plants, such as cushion plants, may be particularly sensitive to climate warming because of their compact growth form and high branch density. In the oceanic southern hemisphere, cushion communities tend to have large range distributions at low latitudes (sea level to low alpine), thus providing an opportunity to test the effects of temperature on plant morphology and reproduction across gradients. Using Donatia novae‐zelandiae as a model species, we compared the leaf morphology, reproduction and responses to warming. Two low‐alpine sites (Maungatua (880 m a.s.l.), Blue Mountains (1000 m a.s.l.)) and two sea‐level sites (Waituna 1 (0 m a.s.l.), Waituna 2 (0 m a.s.l.)) in South Island, New Zealand were used. Donatia novae‐zelandiae cushions differed significantly between the high‐elevation and sea‐level sites both morphologically and in terms of reproduction. High‐elevation cushions produced more flowers (threefold more flowers per plant) and seeds (sevenfold more seeds per capsule) than at sea level, but leaves were larger at sea level (in length and specific leaf area). The cushions were also twice as compact at the high‐elevation sites. After two growing seasons of artificial warming, seed production (35%), leaf length (7%) and width (13%), and specific leaf area (63%) significantly decreased in D. novae‐zelandiae plants; flower production was not significantly affected. Cushion plant morphology and reproduction were significantly affected by environmental drivers at their establishment sites, but all populations responded negatively to artificial warming of 1–3°C. Many cushion plants are considered keystone species because of their propensity to facilitate the growth and establishment of other plant species, the inferred negative effects of global warming on cushion plant species may have a cascading effect on other alpine plant groups.     

Species-specific flowering phenology responses to experimental warming and drought alter herbaceous plant species overlap in a temperate–boreal forest community

Background and AimsWarmer temperatures and altered precipitation patterns are expected to continue to occur as the climate changes. How these changes will impact the flowering phenology of herbaceous perennials in northern forests is poorly understood but could have consequences for forest functioning and species interactions. Here, we examine the flowering phenology responses of five herbaceous perennials to experimental warming and reduced summer rainfall over 3 years.MethodsThis study is part of the B4WarmED experiment located at two sites in northern Minnesota, USA. Three levels of warming (ambient, +1.6 °C and +3.1 °C) were crossed with two rainfall manipulations (ambient and 27 % reduced growing season rainfall).Key ResultsWe observed species-specific responses to the experimental treatments. Warming alone advanced flowering for four species. Most notably, the two autumn blooming species showed the strongest advance of flowering to warming. Reduced rainfall alone advanced flowering for one autumn blooming species and delayed flowering for the other, with no significant impact on the three early blooming species. Only one species, Solidago spp., showed an interactive response to warming and rainfall manipulation by advancing in +1.6 °C warming (regardless of rainfall manipulation) but not advancing in the warmest, driest treatment. Species-specific responses led to changes in temporal overlap between species. Most notably, the two autumn blooming species diverged significantly in their flowering timing. In ambient conditions, these two species flowered within the same week. In the warmest, driest treatment, flowering occurred over a month apart.ConclusionsHerbaceous species may differ in how they respond to future climate conditions. Changes to phenology may lead to fewer resources for insects or a mismatch between plants and pollinators.     

滇西北高原典型湿地湖滨带优势植物种子繁殖对增温的响应

基于IPCC对未来大气增温的预测,采用开顶式生长室(OTC)增温技术,以不增温为对照,研究滇西北高原典型湿地纳帕海湖滨带优势植物水葱和黑三棱在大气增温2.0和3.5℃处理下种子的繁殖特征.结果表明:大气增温对植物结实率影响显著,但存在种间差异.其中,增温对水葱的结实率有促进作用;黑三棱结实率在增温2.0℃处理下显著下降,但在增温3.5℃处理下又恢复到对照水平.增温促进了2种植物穗的生长,在增温2.0、3.5℃处理下水葱的穗长分别增加82.9%、89.0%,小穗数分别增加133.3%、150.0%,每株穗生物量分别增加了10.1%、89.6%,每株穗生物量占总生物量的比例分别增加79.5%、409.3%;在增温2.0、3.5℃处理下,黑三棱穗长分别增加66.1%、95.2%,每株穗生物量占总生物量的比例分别增加878.8%、1052.6%.增温显著增加了水葱和黑三棱每穗种子产量,在增温2.0、3.5℃处理下,水葱每穗种子产量分别增加33.7%、58.3%,黑三棱每穗种子产量分别显著增加3.4%、69.5%.在增温2.0、3.5℃处理下水葱种子长分别增加5.4%、6.9%,种子长宽比分别增大9.1%、5.3%;而增温对黑三棱的种子形态无显著影响.最高温和最低温是影响2种植物种子繁殖的主要温度因子.温度增高引起的生长季提前、营养生长期延长以及有机物积累量增加,为植物繁殖扩散提供了充足的物质和能量积累,可能是2种植物在增温条件下繁殖能力提高的原因.     

Responses to natural climatic variation and experimental warming in two tundra plant species with contrasting life forms: Cassiope tetragona and Ranunculus nivalis

Two circumpolar tundra plant species, the evergreen dwarfshrub Cassiope tetragona and the perennial herb Ranunculus nivalis , were studied at Latnjajaure in northern Swedish Lapland during three consecutive growing seasons (1993–95) as a contribution to the ITEX programme. Open-top chambers (OTCs) were used in a passive heating experiment, and the performance of the plants in unmanipulated controls was correlated with climatic fluctuations among the years. Phenological, vegetative, and reproductive variables were measured. In both species phenological responses were controlled mainly by ambient air temperature. In the evergreen C. tetragona vegetative growth was controlled mainly by the influx of global solar radiation and was not temperature-dependent, whereas the opposite applied in the herbaceous R. nivalis . Vegetative growth in C. tetragona was rather stable among years as well as between treatments, whereas it was strongly influenced by annual climate in R. nivalis . Both species increased their reproductive success with increasing temperature, but R. nivalis was also radiation-dependent in this case, probably because of its green, photosynthetic nutlets. Ovule number in R. nivalis increased steadily in the experimentally heated plots during the study in response to the constant temperature amelioration above the ambient. At the community level, evergreen C. tetragona seems to have low competitive ability under warmer conditions. The situation for vernal low-growing herbs like R. nivalis is more complex; despite a strong positive response to increased temperature, they may exhibit decreased reproductive success if overgrown by a vigorous graminoid canopy.     

Seasonal changes in fluxes of methane and volatile sulfur compounds from rice paddies and their concentrations in soil water

Rice paddies emit not only methane but also several volatile sulfur compounds such as dimethyl sulfide (DMS: CH₃SCH₃). However, little is known about DMS emission from rice paddies. Fluxes of methane and DMS, and the concentrations of methane and several volatile sulfur compounds including hydrogen sulfide (H₂S), carbonyl disulfide (CS₂), methyl mercaptan (CH₃SH) and DMS in soil water and flood water were measured in four lysimeter rice paddies (2.5 × 4 m, depth 2.0 m) once per week throughout the entire cultivation period in 1995 in Tsukuba, Japan. The addition of exogenous organic matter (rice straw) was also examined for its influence on methane or DMS emissions. Methane fluxes greatly differed between treatments in which rice straw had been incorporated into the paddy soil (rice straw plot) and plots without rice straw (mineral fertilizer plot). The annual methane emission from the rice straw plots (37.7 g m^-2) was approximately 8 times higher than that from the mineral fertilizer plots (4.8 g m^-2). Application of rice straw had little influence on DMS fluxes. Significant diurnal and seasonal changes in DMS fluxes were observed. Peak DMS fluxes were found around noon. DMS was emitted from the flood water in the early growth stage of rice and began to be emitted from rice plants during the middle stage. DMS fluxes increased with the growth of rice plants and the highest flux, 15.1 µg m^-2 h^-1, was recorded before heading. DMS in the soil water was negligible during the entire cultivation period. These facts indicate that the DMS emitted from rice paddies is produced by metabolic processes in rice plants. The total amount of DMS emitted from rice paddies over the cultivated period was estimated to be approximately 5–6 mg m^-2. CH₃SH was emitted only from flood water during the first month after flooding.