Hierarchical large-scale to local-scale influence of abiotic factors in summer-fragmented Mediterranean rivers: structuring effects on fish distributions,assemblage composition and species richness

Summer rainfall shortage and low or no flows in Mediterranean rivers make it difficult for fish to attain their final preferenda and hinder the assembling of fish communities. We investigated in this study the degree to which large-scale features of the watersheds and reach-scale habitats correlated with a non-random structure of fish species' richness, composition and distribution under these conditions by intensively sampling a river system during summer and using Geographical Information Systems data. The results of multivariate ordination and association analyses showed that multi-scale factors were hierarchically related and that they correlated with non-random fish distributions and assemblage structure. They also highlighted large-scale factors shaping reach-scale water availability for fish during summer. Fish species' richness, composition and distribution varied along a gradient of water shortage and connectivity loss which were more severe with the smaller drainage area into the reaches upstream. Fish species' distributions along these gradients were strongly correlated with both life-history traits (adult size) and biogeographical origin (native vs. exotic). The results for the summer environmental segregation of exotic versus small endemic richness characteristics indicate strong species' interactions forcing the latter into the upstream, harsher seasonal habitats. Special fitness costs are to be expected in drought years, in regulated rivers, and in general, in a climate change scenario in Mediterranean river networks. The study emphasizes that the conservation of the threatened Mediterranean freshwater fish faunas requires operating at the proper spatial scale. In particular, reach-scale habitat improvements are adequate but not enough during the summer in fragmenting river networks, unless there is proper watershed-scale management of the flow regimes.     

The influence of freshwater flows on two estuarine resident fish species show differential sensitivity to the impacts of drought,flood and climate change

Estuaries are particularly susceptible to climate change and drought resulting in atypical changes to freshwater flows. How such changes in flow impact on the ecology of estuarine fishes may depend on how a species moves in response to changing flow conditions. Acoustic telemetry was used to interpret fine-scale movements of two co-inhabiting estuarine fish species, black bream, Acanthopagrus butcheri and estuary perch, Macquaria colonorum in relation to freshwater flows, season and moon phase. We found black bream to be highly mobile, regularly travelling the length of the estuary and into the neighbouring estuaries. In contrast, estuary perch had particular home ranges and made occasional, upstream or downstream movements. Possibly influenced by freshwater flows, estuary perch moved at greater rates in the Tambo compared to fish in the Mitchell. Black bream resided in the upper estuary during winter and spring and the lower estuary during summer and autumn, whereas estuary perch remained in the upper estuary throughout the year, with occasional downstream movements in winter and spring. This study revealed 1) significantly large increases in freshwater flows result in mass downstream movements in both species, 2) fish moved upstream during full moons and 3) there are contrasting spatio-temporal patterns in movement between species. The results from this study highlight that estuarine fishes are likely to show differential sensitivity to the impacts of drought and climate change and illustrate how acoustic telemetry methods can be used to determine the environmental needs of fishes and help efforts to conserve and manage estuaries worldwide.     

Pesticides and Ostreid Herpesvirus 1 Infection in the Pacific Oyster,Crassostrea gigas

Since 2008, mass mortality outbreaks have been reported in all French regions producing Pacific oysters, and in several Member States of the European Union. These mass mortality events of Pacific oysters are related to OsHV-1 infection. They occur during spring and summer periods leaving suspect the quality of the marine environment and the role of seasonal use of pesticides associated with the arrival of freshwater in oyster rearing areas. Pesticides have been also detected in French coastal waters, especially in areas of oyster production. Using PMA real-time PCR we showed that a mixture of 14 pesticides has no effect on the integrity of virus capsids from viral suspension in the conditions tested. A contact of oysters with this pesticide mixture was related to higher mortality rates among experimentally infected animals in comparison with control ones (no previous pesticide exposure before experimental infection). We therefore suggest that pesticides at realistic concentration can exert adverse effects on Pacific oysters and causes an increased susceptibility to the viral infection in experimental conditions.     

我国南方红壤区氮磷湿沉降对森林流域氮磷输出及水质的影响

持续高通量的氮、磷输入导致水体富营养化的问题已引起广泛关注。通过对江西省千烟洲香溪流域水样(常规水样,降雨后的地表径流以及雨水水样)的季节性监测,研究大气氮、磷湿沉降对森林流域氮、磷输出动态及水质的影响。结果表明:从2013年6月至2014年5月,香溪流域内氮、磷湿沉降通量分别为11.86 kg/hm~2和0.38 kg/hm~2,其中氮湿沉降主要集中在夏秋两季,占全年输入量的64%,而磷沉降主要集中在夏季,占全年输入量的43%,表现出明显的季节性差异;水体p H值(6.22—8.89)的变化范围较大,而且氮、磷的输出受土地管理(施肥方式)及降雨事件的影响较为明显,尤其在耕作期,总氮的输出量占全年氮输出总量的96.2%,而总磷的输出量占全年磷输出总量的61.4%;对4场不同强度降雨(按降雨强度从大到小)的氮、磷输出动态过程分析,发现不同强度的降雨对水体氮、磷的输出过程影响不同,在径流未形成前以及降雨强度达到暴雨级别时,降雨对流域水体氮、磷的稀释作用明显,而在大雨强度下水体磷的输出量明显高于其他降雨;研究期间,香溪流域内氮湿沉降对水体的贡献量为101.97 kg,磷湿沉降的贡献量为0.60 kg,4场降雨氮对流域水体的贡献量为4.46kg,占流域氮输出负荷的15.22%,磷对水体的贡献量为0.032kg,占流域磷输出负荷的0.85%。同时,根据营养状态指数(EI),发现流域水体全年处于中至富营养状态,而且研究期间水体氮、磷浓度均超过水体富营养化阈值(氮1.5 mg/L,磷0.15 mg/L),存在爆发水体富营养化的威胁。     

Species-mediated soil moisture availability and patchy establishment of Pseudotsuga menziesii in chaparral

The occurrence of mature individuals of Pseudotsuga menziesii in stands of Arctostaphylos species mark the initial stages of mixed evergreen forest invasion into chaparral in central coastal California. We planted two cohorts of P. menziesii seeds at three sites under stands of two Arctostaphylos species and Adenostoma fasciculatum in order to determine whether first-year seedling emergence and survival, particularly during the regular summer drought, underlie the spatial distribution of mature trees observed in chaparral. Regardless of the chaparral species they were planted under, P. menziesii seeds that were not protected from vertebrate predation displayed very little emergence and no survival. In contrast, emergence of P. menziesii that were protected from vertebrate predators was much higher but still did not significantly differ among the three chaparral species. However, survival of protected seedlings under Arctostaphylos glandulosa was much greater than under A. fasciculatum, with intermediate survival under Arctostaphylos montana. While mortality of protected seedlings due to insect herbivory, fungal infection, and disturbance displayed no consistent patterns, summer drought mortality appeared to drive the patterns of survival of P. menziesii under the different chaparral species. These emergence, mortality, and survival data suggest that spatial patterns of P. menziesii recruitment in chaparral are driven by first-year summer drought seedling mortality, but only in years when seeds and seedlings are released from vertebrate predation pressure. Because the first-year drought mortality and survival patterns of P. menziesii seedlings differed strongly depending on the chaparral species, we examined the additional hypothesis that these patterns are associated with differences in the availability of soil moisture under different chaparral species. Both higher survival and lower drought mortality of P. menziesii seedlings were associated with higher soil water potential under Arctostaphylos stands during the summer drought, especially in the subsurface soil. The data suggest that Arctostaphylos stands, particularly stands of A. glandulosa, ameliorate xeric summer conditions to a degree that facilitates first-year establishment of P. menziesii and strongly influences spatial distribution of mature trees. Received: 18 September 1998 / Accepted: 23 December 1998     

Synergistic impacts of heat shock and spawning on the physiology and immune health of Crassostrea gigas: an explanation for summer mortality in Pacific oysters

Mass mortality is often observed in cultured oysters during the period following spawning in the summer season. To examine the underlying causes leading to this phenomenon, thermotolerance of the Pacific oyster Crassostrea gigas was assessed using pre- and postspawning oysters that were sequentially treated with sublethal (37 degrees C) and lethal heat shocks (44 degrees C). The effects were examined on a range of immune and metabolic parameters in addition to mortality rate. A preventative 37 degrees C significantly reduced oyster mortality after exposure to a second heat shock of 44 degrees C, but in postspawning oysters mortality remained at 80%, compared with < 10% in prespawning oysters. Levels of the 72 kDa and 69 kDa heat shock proteins were low in the gill tissue from postspawning oysters stimulated by heat shock, indicating spawning reduced heat shock protein synthesis. The postspawning oysters had depleted glycogen stores in the mantle tissue and reduced adenylate energy charge after heat shock, indicative of lower energy for metabolic activity. A cumulative effect of spawning and heat shock was observed on the immunocompetence of oysters, demonstrated by reduced hemocyte phagocytosis and hemolymph antimicrobial activity. These results support the hypothesis that the energy expended during reproduction compromises the thermotolerance and immune status of oysters, leaving them easily subject to mortality if heat stress occurs in postspawning stage. This study improves our understanding of oyster summer mortality and has implications for the long-term persistence of mollusks under the influence of global warming.     

Host-dependent genetic variation in freshwater pearl mussel (Margaritifera margaritifera L.)

Freshwater fishes are vulnerable to changes in water quality, physical habitat and connectivity resulting from drought, particularly in regulated rivers. When adequate river flows return, the recovery of populations might depend on the duration and consequences of drought. Rivers of the highly regulated Murray–Darling Basin in south-eastern Australia terminate at two large, shallow lakes that are separated from the estuary by tidal barrages. Over-abstraction of water and widespread prolonged drought (1997–2010) placed the lakes under severe environmental stress, culminating in critical water level recession from 2007 to 2010. Concurrently, most freshwater fish populations collapsed. We investigate shifts in fish assemblages resulting from habitat inundation in the lakes following the drought. The inundation and re-connection of the lakes and fringing habitats led to a substantial reduction of salinity throughout the region, and aquatic vegetation shifted from salt-tolerant to freshwater species. Fish assemblages became increasingly characterized by common freshwater taxa (ecological generalists), including high proportions of alien species. There were no indications of population recovery for three threatened species. The findings emphasize that short-lived fishes with specialized habitat requirements are vulnerable to severe population declines during prolonged drought in regulated rivers, which might restrict their recovery when adequate flows return.     

Experimental manipulations of water and nutrient input to a Norway spruce plantation at Klosterhede,Denmark

At Klosterhede, Denmark experimental manipulations of water and nutrient inputs to a forest stand were carried out under a 1200 m² roof simulating i) summer drought, ii) removal of N and S input, and iii) optimal nutrition and water availability. In practise, manipulation of element fluxes in a complex system like a forest may cause unintended changes such as climatic effects from the roof which may interfere with the results of the intended manipulation. This paper illustrates the extent and effects of such unintended changes.The most important climatic change under the roof was a 50% reduction of photosynthetic light which caused a decrease in moss cover on the ground floor. Soil temperatures under the roof were decreased 0.3°C during summer and increased 0.2°C during winter. Air humidity was not changed. Throughfall water amount decreased close to the stems whereas element concentrations in throughfall increased close to the stems and with tree size on the ambient plots. This distinct spatial pattern was changed under the roof to a more variable water distribution from the sprinkling system and to constant element concentrations in the sprinkling water. This loss/change of spatial variability in the input was reflected in the soil solution. The concentration patterns found in soil water (increasing close to the stem and with tree size) in the ambient plots were no longer observed under the roof.Exclusion of throughfall by the roof disturbed the internal cycle of nutrients leached from the canopy. The removal of canopy leached Ca and K by the roof caused a decline in soil water concentrations, even on the drought plot where natural throughfall was only cut off during two summer months. The internal cycle of Ca and K had to be restored by addition of Ca and K under the roof. Further, leachable elements and soluble compounds may be washed out from litterfall during the collection period on the roof, and thus lost from the internal cycle by by exclusion of throughfall.It is important to consider these problems in the design of new manipulation experiments and in the interpretation of the results. Some recommendations are given.     

Effects of freshwater availability on the summer distribution of otters Lutra lutra in the southwest coast of Portugal

The availability of freshwater has been suggested to strongly influence the distribution of coastal otters Lutra lutra The test this hypothesis, a study was undertaken on the relationships between otter distribution and freshwater availability in a coast where freshwater is a very scarce resource during the summer In this area otters occur mostly in a few small coastal streams but feed largely in the sea Twentynine streams were surveyed in the summer of 1990, and the presence/absence of otter signs was related to nine habitat variables Freshwater availability was found to be the most important factor influencing the occurrence of otters in summer It is suggested that otters favour large streams with good vegetation cover, for these are the most likely to maintain frestwater during the dry periods The distribution of otter signs m streams was also analysed, and it was found that signs tend to concentrate particularly near the mouth of the streams This habitat analysis has some conservation implications, indicating that the decrease of freshwater availability or quality of streams can make these unsuitable for otters, severely decreasing the areas suitable for the species in the southwest coast of Portugal     

Effects of Climate Variability and Accelerated Forest Thinning on Watershed-Scale Runoff in Southwestern USA Ponderosa Pine Forests

The recent mortality of up to 20% of forests and woodlands in the southwestern United States, along with declining stream flows and projected future water shortages, heightens the need to understand how management practices can enhance forest resilience and functioning under unprecedented scales of drought and wildfire. To address this challenge, a combination of mechanical thinning and fire treatments are planned for 238,000 hectares (588,000 acres) of ponderosa pine (Pinus ponderosa) forests across central Arizona, USA. Mechanical thinning can increase runoff at fine scales, as well as reduce fire risk and tree water stress during drought, but the effects of this practice have not been studied at scales commensurate with recent forest disturbances or under a highly variable climate. Modifying a historical runoff model, we constructed scenarios to estimate increases in runoff from thinning ponderosa pine at the landscape and watershed scales based on driving variables: pace, extent and intensity of forest treatments and variability in winter precipitation. We found that runoff on thinned forests was about 20% greater than unthinned forests, regardless of whether treatments occurred in a drought or pluvial period. The magnitude of this increase is similar to observed declines in snowpack for the region, suggesting that accelerated thinning may lessen runoff losses due to warming effects. Gains in runoff were temporary (six years after treatment) and modest when compared to mean annual runoff from the study watersheds (0–3%). Nonetheless gains observed during drought periods could play a role in augmenting river flows on a seasonal basis, improving conditions for water-dependent natural resources, as well as benefit water supplies for downstream communities. Results of this study and others suggest that accelerated forest thinning at large scales could improve the water balance and resilience of forests and sustain the ecosystem services they provide.     

Changes in the association between summer temperature and mortality in Seoul, South Korea

The health impact of climate change depends on various conditions at any given time and place, as well as on the person. Temporal variations in the relationship between high temperature and mortality need to be explored in depth to explain how changes in the level of exposure and public health interventions modify the temperature–mortality relationship. We examined changes in the relationship between human mortality and temperature in Seoul, which has the highest population in South Korea, considering the change in population structure from 1993–2009. Poisson regression models were used to estimate short-term temperature-related mortality impacts. Temperature-related risks were divided into two “time periods” of approximately equal length (1993 and 1995–2000, and 2001–2009), and were also examined according to early summer and late summer. Temperature-related mortality in summer over the past 17 years has declined. These decreasing patterns were stronger for cardiovascular disease-related mortality than for all non-accidental deaths. The novel finding is that declines in temperature-related mortality were particularly noteworthy in late summer. Our results indicate that temperature-related mortality is decreasing in Seoul, particularly during late summer and, to a lesser extent, during early summer. This information would be useful for detailed public health preparedness for hot weather.     

Survival and recovery of perennial forage grasses under prolonged Mediterranean drought: I. Growth, death, water relations and solute content in herbage and stubble

Swards of Dactylis glomerata cultivars (cvs) KM2 and Lutetia and of Lolium perenne cvs Aurora and Vigor were grown under full irrigation or prolonged summer drought (80 d) in a field experiment in the South of France.
After irrigation was withheld, leaf extension rates of all cvs fell by 90% within 9–12 d, and rapid scorching of laminae followed. Tiller mortality at the end of the drought was very different in the cocksfoot cvs (4% for KM2 and 76% for Lutetia) and intermediate (41%) for both ryegrass cvs. Following re-watering, rates of herbage regrowth were closely correlated with tiller survival. Measured minerals contributed c . 0·52 MPa to osmotic potential in all treatments, whereas water-soluble carbohydrates (WSC) contributed 0·25 MPa under irrigation and 0·46 MPa during drought.
There was no systematic difference between the two species for summer survival under severe drought, but large differences between the cocksfoot cvs. The traits most strongly associated with superior survival were: (a) a deep root system and greater water uptake at depth; (b) low water and osmotic potentials in surviving laminae, i.e. better tolerance to dehydration; (c) large pool-size of WSC reserves (fructans having degree of polymerization >4) in entire tiller bases (stubble); (d) low accumulation of proline in stubble; (e) rapid nitrogen uptake after rewatering.     

Mass mortality in Pacific oysters is associated with a specific gene expression signature

Mass mortality events occur in natural and cultured communities of bivalve molluscs. The Pacific oyster, Crassostrea gigas, is a dominant species in many intertidal locations as well as an important aquacultured bivalve species, and for the last 50 years, adult oysters have suffered frequent and extreme mass mortality events during summer months. To investigate the molecular changes that precede these mortality events, we employed a novel nonlethal sampling approach to collect haemolymph samples from individual oysters during the period that preceded a mortality event. Microarray-based gene expression screening of the collected haemolymph was used to identify a mortality gene expression signature that distinguished oysters that survived the mortality event from those individuals that died during the event. The signature was cross-validated by comparing two separate episodes of mortality. Here, we report that near-mortality oysters can be distinguished from longer-lived oysters by the elevated expression of genes associated with cell death, lysosomal proteolysis, and cellular assembly and organization. These results show the potential utility of nonlethal sampling approaches for investigating the environmental causes of mortality in natural populations in the field, and for predicting when such events could occur and which individuals will be affected.     

Genetic polymorphism of glutamine synthetase and delta-9 desaturase in families of Pacific oyster Crassostrea gigas and susceptibility to summer mortality

Large-scale mortality events have been observed in Pacific oyster Crassostrea gigas on the west coast of France since the early 1980s, particularly during summer. In order to understand the causes of this mortality, two generations of oysters from single-pair matings were studied in three sites on the French Atlantic coast (Baie-des-Veys, Auray and Ronce-les-Bains). The present paper examines the role of two candidate genes in the susceptibility of oysters to summer mortality, and the selective pressure exerted by such mortality on their polymorphism. Glutamine synthetase (amino-acid metabolism) and delta-9 desaturase (lipid metabolism) genes were studied in the successive generations, using polymerase chain reaction single-strand conformation polymorphism (PCR-SSCP). Observed and expected genotypic frequencies were compared. Three different alleles were detected for the glutamine synthetase gene and two for delta-9 desaturase. Allele C of glutamine synthetase seemed to be counter-selected in some second generation families. Allele B of delta-9 desaturase gene was potentially counter-selected at Auray in the families showing higher mortality, and strong selection against BB homozygotes was observed. These observations led us to conclude that any selective effect of summer mortality on allele C of glutamine synthetase gene or allele B of delta-9 desaturase gene could be mediated either directly or via linkage to other loci involved in physiological pathways affecting susceptibility.     

Recovery from Severe Mistletoe Infection After Heat- and Drought-Induced Mistletoe Death

Ecosystems - Mistletoes are emerging as important co-contributors to tree mortality across terrestrial ecosystems, particularly when infected trees are stressed by water limitations during drought....     

Water conservation in<Emphasis Type="Italic"> Artemisia tridentata</Emphasis> through redistribution of precipitation

Water conservation is important for plants that maintain physiologically active foliage during prolonged periods of drought. A variety of mechanisms for water conservation exist including stomatal regulation, foliage loss, above- and below-ground allocation patterns, size of xylem vessels and leaf pubescence. Using the results of a field and simulation study with Artemisia tridentata in the Great Basin, USA, we propose an additional mechanism of water conservation that can be used by plants in arid and semi-arid environments following pulses of water availability. Precipitation redistributed more uniformly in the soil column by roots (hydraulic redistribution of water downward) slows the rate at which this water can subsequently be taken up by plants, thus prolonging water availability during periods of drought. By spreading out water more uniformly in the soil column at lower water potentials following precipitation events, water use is reduced due to lower soil conductivity. The greater remaining soil water and more uniform distribution result in higher plant predawn water potentials and transpiration rates later in the drought period. Simulation results indicate that plants can benefit during drought periods from water storage following both summer rain events (small summer pulses) and overwinter recharge (large spring pulse). This mechanism of water conservation may aid in sustaining active foliage, maintaining root-soil hydraulic connectivity, and increasing survival probability of plants which remain physiologically active during periods of drought.     

Streams in an uninhabited watershed have predictably different thermal sensitivities to variable summer air temperatures

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Prevalence and infection intensity of the ovarian parasite Marteilioides chungmuensis during an annual reproductive cycle of the oyster Crassostrea gigas

The occurrence of Marteilioides chungmuensis, a protozoan paramyxean parasite in the reproductive system of the Pacific oyster Crassostrea gigas, was observed at Gosung Bay, Korea. Seasonal variation in gonad development was investigated in a suspended cultured oyster population. Gametogenesis began in February and first-spawning was observed between mid and late June when surface water temperature reached 22 to 25 degrees C. Spawning activity extended from mid June to late September, with 2 marked spawning peaks in June and August. Histological examination indicated that gonad development paralleled seasonal fluctuations in water temperature. Spawning in late June was partly associated with a sudden drop in salinity due to large freshwater inputs to the Bay with the summer monsoon. M. chungmuensis occurred in developing and fully mature eggs of spawning oysters in late June to January, but were not observed from February to May. Monthly mean infection intensity was high in late June when most oysters had their first spawning period. The infection level was also relatively high in late August and November, when oysters were spawning or had completed spawning. Several oysters collected in November (11.4%) and December (16.3%) carried a large quantity of ripe but M. chungmuensis-infected eggs, suggesting that infection also causes spawning failure by delaying spawning and destroying ripe oocytes.     

Responses of a CAM Plant to Drought and Rainfall: Capacitance and Osmotic Pressure Influences on Water Movement

Daily and seasonal patterns in water flow and water potentialwere investigated for the Crassulacean acid metabolism succulentAgave deserti during an extended summer drought and for a periodfollowing rainfall. Field measurements of transpiration andof osmotic pressure changes over selected 24 h periods wereused as input variables for a computer model of water flow thatwas based on an electrical circuit analog of the whole plant.Parameters such as root resistance and tissue capacitance werealso varied to reflect the effects of changing plant or soilwater status. The model predicted internal water flow and waterpotential during the drought cycle and was used to assess therole of tissue osmotic properties in water uptake from the soiland in internal water redistribution. For plants under wet soil conditions, 55% of the night-timetranspiration was derived from water storage, this storage beingrecharged during the day. As drought progressed, transpirationand the nocturnal increase in osmotic pressure declined, althoughthe osmotic pressure itself increased. The difference in osmoticpressure between the water storage tissue and the chlorenchymacaused a net flow of water into the chlorenchyma after 3 weeksof drought, thereby increasing chlorenchyma turgor pressure.Simulations also indicated that a large increase in root resistancemust occur to prevent substantial water loss from the plantto the dry soil. After rainfall, recharge of plant water storagewas complete within one week, although full recovery in theamplitude of daily osmotic pressure variations took longer. Key words: Agave deserti, transpiration, water potential, water storage     

干旱胁迫下白刺花种子大小与萌发对策

种子大小与种子萌发及其与环境因子的关系是植物种子萌发对策研究中的重要科学问题之一。采用PEG模拟干旱法研究不同干旱胁迫强度(0,5%,10%,15%,20%)下,白刺花(Sophora davidii)种子萌发进程、种子大小与种子萌发及种子命运的关系。结果表明:不同干旱胁迫下,白刺花种子具有相似的萌发进程,但中度干旱处理(10%PEG)萌发率显著高于零干旱(0%PEG)和重度干旱处理(P0.05),重度干旱处理(20%PEG)种子萌发开始时间晚于零干旱和中度干旱处理;种子大小与种子萌发开始时间的关系表现为零干旱处理下呈极显著负线性关系,中度干旱处理(5%PEG,10%PEG)下无相关关系,重度干旱处理(15%PEG,20%PEG)下呈负二次曲线关系;种子大小对种子命运的影响表现为零干旱处理有利于大、小种子萌发和小种子休眠,中度干旱处理(10%PEG)增加中等种子萌发、大种子休眠和小种子死亡风险,重度干旱处理(15%PEG,20%PEG)增加大种子死亡风险、中等种子和小种子休眠。综合分析表明,白刺花种子大小与萌发行为及种子命运的关系具有较强的环境依赖性,即种子萌发行为表现为顺境下种子越大萌发越快,逆境下小种子和大种子较中等种子萌发更快;种子命运表现为顺境增加种子死亡的风险,中度干扰有利于种子萌发,逆境则有利于种子休眠。