Population dynamics of zooxanthellae during a bacterial bleaching event

Each summer 80–90% of the colonies of Oculina patagonica undergo bleaching off the Mediterranean coast of Israel. To investigate fluctuations through a yearly bleaching cycle, monthly measurements of zooxanthella density, mitotic index and chlorophyll-a concentration were conducted. Results showed (1) a significant negative correlation between sea surface temperature (SST) and zooxanthella density; (2) both significantly lower zooxanthella mitotic index and higher chlorophyll-a per zooxanthella content during the bleaching season compared with the non-bleaching period; (3) prior to bleaching, a lag between the peak of zooxanthella density and chlorophyll-a concentration followed by a similar lag during recovery. Zooxanthella density declined significantly between March and May while chlorophyll-a concentration peaked in April, and then declined. Zooxanthella density increased significantly in November while chlorophyll-a concentration increased significantly in January. We conclude that during bacterial bleaching events, zooxanthellae are severely damaged. However, by the time of the following bleaching event the coral tissues regain their “normal” (pre-bleaching) zooxanthella population density.     

Distribution and movement of toxaphene in anaerobic saline marsh soils

The biological oxygen demand (BOD) of filtered water from Lake Wingra, Wisconsin is significantly higher in the littoral zone than in the pelagial zone. Laboratory experiments indicate that BOD is not influenced by water temperature at the time of sampling or by enrichment with nitrate or ammonia. Rather, enrichment with macrophyte leachate sharply increases BOD, and enrichment with phosphate produces a small but significant increase in BOD. We conclude that high BOD in littoral waters of the lake is an indication of production of labile organic matter in the water by dense beds of the macrophyte Myriophyllum spicatum.     

A flow injection analysis system with encapsulated high-density <Emphasis Type="Italic">Saccharomyces cerevisiae</Emphasis> cells for rapid determination of biochemical oxygen demand

The biochemical oxygen demand (BOD) determination was studied using a novel flow injection analysis (FIA) system with encapsulated Saccharomyces cerevisiae cells and an oxygen electrode and was compared with conventional 5-day BOD tests. S. cerevisiae cells were packed in a calcium alginate capsule at a dry cell weight of 250 g/l of capsule core. The level of dissolved oxygen (DO) was reduced due to the enhanced respiratory activity of the microbial cells when the injected nutrient passed through the bioreactor. The decrease in DO (ΔDO) was intensified with the amount of microbial cells packed in the bioreactor. However, the specific ΔDO decreased as the amount of cells loaded in the bioreactor increased. The ΔDO value was dependent on the pH and temperature of the mobile phase and reached its maximum value at 35°C and pH 7–8. Also, ΔDO became larger at longer response times as the flow rate of the mobile phase decreased. The measurement of ΔDO was repeated more than six times consecutively using a 20-ppm standard glucose and glutamic acid solution, which confirmed the reproducibility with a standard deviation of 0.95%. A strong linear correlation between ΔDO and BOD was also observed. The 5-day BOD values of actual water and wastewater samples were in accordance with the BOD values obtained by this FIA method using encapsulated S. cerevisiae cells. Unlike the cell-immobilized bead system, there was no contamination of the bioreactor resulting from any leak of yeast cells from the sensor capsules during BOD measurements.     

Does the Number of Diapausing Eggs Laid on Bark Affect the Population Dynamics of the Spider Mite Panonychus mori?

We studied the relationship between number of diapausing eggs produced by the spider mite Panonychus mori and the subsequent population trend for a period of 3 years. Panonychus mori showed a single population density peak on its host plant moonseed, Cocculus trilobus. The position and height of this peak were correlated with the density of diapausing eggs around the moonseed leaf buds produced the winter before. In 1994 the density of diapausing eggs measured in February was 4.3/bud, which was 6–14 times higher than the density of diapausing eggs for the same period in 1995 (0.3/bud) and 1996 (0.7/bud). The subsequent population density peak in 1994 occurred in mid June and was about 2.5 times higher than the peaks in 1995 and 1996, which both occurred early September. Thus, the present study showed a positive correlation between the density of diapausing eggs on the host plant and the start and the extend of the population increase the next growing season. Predators associated with the spider mite population were phytoseiid mites, especially Amblyseius eharai was well synchronized with the spider mite density in 1994. Field observations revealed that P. mori produced diapausing eggs in response to short photoperiod in early October each year, which corresponded with the timing predicted by the critical photoperiod around 13 h at 18°C, as assessed in laboratory trials. Diapause ended by early April when egg hatchability attained about 50% and eggs took 9 days to hatch at 25°C and a 16L:8D photoperiod. Hatching in early April was twice faster than in late February.     

Swimming Behaviour of <Emphasis Type="Italic">Chironomus acerbiphilus</Emphasis> Larvae in Lake Katanuma

We conducted a seasonal survey of the swimming behaviour of Chironomus acerbiphilus larvae in volcanic Lake Katanuma from April 1998 to December 2001. Swimming C. acerbiphilus density was much higher than other chironomid species in lakes. All C. acerbiphilus larvae (1st through 4th instars) swam, but the earlier instars (especially the 1st) had the greatest densities and fluctuations. First instars were never found in the benthic population. This result indicates that the 1st-instar larvae are planktonic. Low water temperature (below about 10 °C) resulted in the seasonal disappearance of swimming chironomid larvae. Chemical factors – oxygen depletion or presence of hydrogen sulfide – also restricted the distribution of swimming and benthic larvae. Larvae were distributed only in the oxygen-rich part of the lake bottom and swam only in the oxygen-rich layer of the water column. The density of older swimming C. acerbiphilus (3rd and 4th instars) tended to increase with increasing benthic larval densities. The chemical stress of oxygen depletion or presence of hydrogen sulfide during holomixis within and after the stratification period leads to conspicuous swimming behaviour of benthic C. acerbiphilus larvae. Almost all C. acerbiphilus larvae died on this occasion.     

Ant predation ofEriocrania miners in a polluted area

We tested the hypothesis that air pollution may affect population densities of birch-feeding leafminers via changes in ant predation. Foraging activity of three ant species (Formica rufa, F. fusca andF. lemani), predation rates and population densities of both solitary and gregariousEriocrania (Lepidoptera: Eriocraniidae) miners were investigated at 13 sites around the Harjavalta copper-nickel smelter, SW Finland. Ant species differed in their distribution patterns relative to pollution. However, the total percentage of birch trees foraged by ants (all species combined) showed no correlation with the distance from the factory complex. As a result, no clear trends in predation rates were apparent in relation to the distance from the pollution source for either solitary or gregariousEriocrania species. Densities of the solitaryEriocrania species tended to increase with the distance from the pollution source whereas densities of the gregariousE. haworthi peaked close to the factory complex. No corresponding differences in predation rates between solitary and gregarious miners were found. Ant predation, thus, did not explain density patterns ofEriocrania miners in the polluted area.     

The carbon monoxide- and oxygen-reacting haemoproteins of Streptomyces clavuligerus: cytochrome aa 3 is the predominant terminal oxidase of the respiratory chain

The nature of the carbon monoxide- and oxygen-reacting haemoproteins in the respiratory chain of the filamentous antibiotic-producing bacterium Streptomyces clavuligerus has been investigated. CO-difference (i.e. CO+ reduced minus reduced) spectra of intact cells showed the presence of cytochrome aa ₃, a CO binding b-type cytochrome, and a pigment resembling cytochrome d. In addition, cells that were approaching the end of the growth phase showed the presence of cytochrome P450: this pigment was undetectable in cells harvested early in the growth cycle. High speed centrifugation of cell-free extracts prepared from cells broken by sonication showed that cytochrome aa ₃ was tightly membrane-bound and that cytochrome P450 was soluble. Inhibition of oxygen uptake rates of cells by cyanide indicated that one component, which showed 50% inhibition at 2–4 mM CN^–, was acting as major terminal oxidase: this was observed in cells harvested from all stages of growth. Photodissociation (i. e. photolysed, CO reduced minus CO reduced) spectra at-118°C, in the absence of oxygen, showed cytochrome aa ₃ to be the sole photolysable CO-reacting haemoprotein. At higher temperature (-87°C), in the presence of oxygen, cytochrome aa ₃ formed a complex with oxygen that could not be photolysed by similar intensities of light. By raising the temperature to-43°C, the oxidation of c-type cytochromes was observed. It is concluded that cytochrome aa ₃ is the predominant terminal oxidase in S. clavuligerus and that the other CO reacting haemoproteins, of unknown function, are unlikely to be oxidases.     

新疆玉龙喀什河与策勒河流域胡杨年轮对气象因子的响应研究

在树木年轮学与气候学理论基础上,以新疆和田地区玉龙喀什河与策勒河下游河岸胡杨为研究对象,基于流域对应的气象、沙尘天气日数、策勒河径流量等因子,分析了玉龙喀什河与策勒河流域胡杨年轮对气象因子的响应特征,以明确干旱荒漠区胡杨生长过程中的气候限制因子,为促进胡杨生态修复及沙尘防治提供理论依据.结果显示:(1)1976-201...     

宝天曼自然保护区不同针叶树径向生长对气候的响应

树木年轮资料能够提供区域内过去长时间的环境和气候信息,成为获取过去气候变化信息的重要手段之一。利用采自宝天曼自然保护区的两种针叶树种油松和华山松树木年轮样本分别建立了油松和华山松树轮宽度标准年表PT和PA,并将油松和华山松样本合并建立了联合树种的区域年表(RC)。3种年表分别与不同气候要素(月平均气温、月平均最高气温、月平均最低气温、月降水量)及其不同月份组合进行相关分析,结果表明,油松年表PT和华山松年表PA都包含较高的气候信息,且都和生长季不同月份温度显著负相关和降水显著正相关。其中,油松和华山松都与当年4月和5月降水显著正相关,油松还与当年5月的平均最高温度和上年11月最低温显著负相关,与当年3月平均最低气温显著正相关;华山松与上年10月和当年4月的平均最高温度显著负相关,与上年12月和当年7月平均最低温度显著正相关;联合年表RC包含了单个年表PT和PA共同的气候信息,与当年4—5月降水和3月最低温显著正相关,与当年7月最低温和4、5月最高温显著负相关。不同年表与生长季(3—8月)内气候要素月份组合的相关分析也表明联合年表RC包含和单物种年表PT、PA相似的气候信息并加强了PT、PA受当年生长季气候变化影响的公共信号。可见,同一地区的多树种联合年表一定程度上能体现出区域性树木生长对气候变化响应的生态生理特征,为同地区建成多树种联合年表来探讨当地区域性气候变化提供了可行性的理论基础和一定的参考作用.     

Seasonal abundance,breeding and sex-structure of populations of Tilapia nilotica (L.) and Labeo niloticus (F.) in the Jebel Aulia Dam area of the White Nile

Seasonal abundance, maturation and sex-structure were studied in populations of T. nilotica and L. niloticus in the Jebel Aulia reservoir and in the White Nile below the dam. L. niloticus showed a single annual breeding cycle from mid-July to December and was most abundant in October in the lake, and in early November in the river. T. nilotica showed two distinct annual seasons of abundance: summer (April to June) and autumn (November to October). Spawning activity was highest in summer (April to June) and autumn (late July to September). The sex-ratio varied inversely with age (size); males were predominant in populations of older fish, perhaps because of their greater endurance to adverse conditions as evidenced by their better survival in hypoxic water.     

1801年以来河南尧山地区油松高温变化及影响机制

基于建立的河南尧山地区油松树轮宽度标准年表,分析了油松径向生长与该地区气温、降水等气候因子之间的关系以及气温升高前后树木生长与平均最高气温间的相关性,结果表明:4—7月平均最高气温与树轮年表的相关系数高达-0.64,是该地区油松径向生长的主要限制因子;气候变暖后树轮年表与9、10月份平均最高气温显著正相关,与2、3月降水显著正相关,与4—7月平均最高气温间的相关性较为稳定。因此,重建了尧山地区1801年来4—7月平均最高气温,其方差解释量达40%(调整自由度后为38.9%)。过去216年的高温重建历史中经历了6个较暖的时期:1801—1825,1845—1853,1876—1889,1922—1944,1957—1975,1996—2013年和5个较冷的时期:1826—1844,1854—1875,1890—1921,1945—1956,1978—1995年,其结果与过去伏牛山龙池曼地区5—7月温度重建序列具有很好的一致性。周期分析结果发现该地区4—7月平均最高气温变化存在着2—4年(ENSO周期)和35.23—48.47年的主要变化周期,小波分析发现在1920年前后气候由长周期变为短周期变化;空间相关分析显示重建的高温序列很好地代表了豫东平原地区的温度变化,同时也发现与北太平洋副热带高压850 hPa上空的温度有非常高的正相关关系,表明豫东高温的波动可能与北太平洋海气振荡有关,这一研究结果为山区森林管理及平原区农业生产提供基础服务数据。     

Effect of rearing temperature and larval density on larval survival, age at pupation and adult size of Anopheles gambiae

The effects of temperature and larval density on survival of larvae, growth rate, age at pupation, and adult size (measured as wing length and dry weight) of laboratory-reared Anopheles gambiae (Diptera: Culicidae) were studied. Larvae were reared at three temperatures (24, 27 and 30°C) and three densities (0.5, 1 and 2 larvae/cm²). The effects of density and temperature strongly interacted to determine the mosquitoes' life-history parameters. Survival was highest at the intermediate temperature of 27°C. The differences between the temperatures increased with increasing density. At 30°C survival decreased as density increased, but at 27°C increasing density led to higher survival. Age at pupation increased as temperature decreased from 30°C to 24°C and as density decreased from 2 to 0.5 larvae/cm². Adult size also increased as temperature decreased, but showed a negative correlation with density only at 27°C. In contrast, at 24°C and 30°C a decrease in density led to a decrease in adult size. Growth rate showed a similar pattern. At 27°C growth rate decreased as density increased, but at other temperatures the opposite trend was observed.     

川西亚高山林线岷江冷杉和红杉对气候变化的响应

为揭示川西亚高山林线优势树种岷江冷杉（Abies fargesii var.faxoniana）和红杉（Larix potaninii）径向生长对气候变化的响应差异。通过树木年轮生态学的方法,研究了岷江冷杉和红杉径向生长与年均温的分异现象、生长衰退以及应对严重干旱事件的抵抗力和恢复力稳定性,并结合限制岷江冷杉和红杉径向生长的主要气候因素进行了分析。结果表明：（1）温度是限制林线岷江冷杉径向生长的主要气候因素,而限制红杉径向生长的主要气候因素是降水;（2）岷江冷杉的径向生长与年平均温度的变化趋势一致,红杉在20世纪90年代后径向生长呈现出下降的趋势,与年均温的升高出现分异;（3）1800-2019年间,岷江冷杉出现生长衰退2次,红杉出现生长衰退5次,相同的时间内,红杉的衰退频率高于岷江冷杉;（4）岷江冷杉对严重干旱事件的恢复力相比升温前有所增强,抵抗力相比升温前下降明显,红杉对严重干旱事件的抵抗力和恢复力相比升温前均呈现出下降的趋势。可为川西亚高山林线的植被动态变化预测和历史区域气候重建提供参考。     

Responses of ring widths and maximum densities of Larix gmelinii to climate on contrasting north- and south-facing slopes in central Siberia

An analysis was performed of the climatic responses of the radial growth of Larix gmelinii (Rupr.) Rupr. at two sites—both of which included contrasting north- and south-facing slopes—in Tura, central Siberia, with the development of ring width and maximum-density chronologies for each slope. Both residual and standard chronologies of ring widths were positively correlated with temperature from late May until mid June on all four slopes. By contrast, standard chronologies of ring widths were negatively correlated with precipitation during the winter (from October to April) and in May on the north-facing slope at site 1 and on the south-facing slope at site 2 respectively. The negative correlations with precipitation during the winter and in May on some of the slopes suggested that delayed snowmelt in early spring might inhibit the radial growth of L. gmelinii, and the effects of snow are likely to vary with topography. Both residual and standard chronologies of maximum densities were positively correlated with temperature in early July on all four slopes. Maximum densities were also positively correlated with precipitation during summer of the previous year on all the slopes. These suggest that no major differences exist in terms of responses of maximum density to climatic factors between the north- and south-facing slopes.     

武夷山不同林龄甜槠林土壤呼吸特征及影响因素

为揭示中亚热带常绿阔叶林群落优势种一甜槠天然林不同林龄林下土壤呼吸(Soil respiration,RS)差异及影响因素,采用LI-8100开路式土壤碳通量系统对武夷山自然保护区不同林龄(18、36、54、72 a)天然甜槠林进行了1年的野外原位测定。结果表明:(1)不同林龄甜槠林RS季节动态呈现明显的单峰趋势,林龄对冬季RS影响并不显著(P>0.05),秋季18 a甜槠林RS与其他3种林龄差异显著(P<0.05),林龄对土壤含水率的季节变化没有显著影响(P>0.05);(2)不同林龄甜槠林5 cm深土壤温度与RS拟合R2明显高于土壤含水率与RS拟合R2,随着林龄增大,RS温度敏感性指数Q10值呈上升趋势,依次为1.551、1.589、1.640、1.664,且54、72 a甜槠林RS温度敏感性指数Q10值显著高于18、36 a(P<0.05);(3)土壤含水率与5 cm深土壤温度共同解释了RS变异的86%—90.3%;0—60 cm土层根系生物量与5 cm深土壤温度共同解释了RS变异的88.3%—91.8%,由此可见,生物因子与非生物因子双因素拟合可以更好地解释不同林龄RS差异。在对未来森林植被土壤呼吸及碳汇功能进行研究时,应在考虑林龄及季节差异的基础上,加强对生物因子的测定。     

野牛草克隆分株酶促活性氧清除系统对差异光周期的响应

以盆栽野牛草克隆分株为材料,将克隆分株分别标记为O(姊株)和Y(妹株),设置连接组和断开组两种处理,其中,连接组中O分株和Y分株通过节间子相连,断开组则剪断分株节间子;两组处理的O分株光周期均设置为光照12h/黑暗12h,Y分株光周期均设置为黑暗12h/12h光照(恰好与O分株相反),经过7d的差异光周期处理后进行72h全光照稳定培养,并于全光照条件下在48h内连续测定各分株叶片超氧化物歧化酶(SOD),过氧化物酶(POD),过氧化氢酶(CAT),抗坏血酸过氧化物酶(APX)的活性以及丙二醛(MDA)的含量,探讨野牛草叶片酶促活性氧清除系统对差异光周期的响应特征。结果表明,差异光周期处理1周后,全光照条件下,断开状态的野牛草克隆分株O和Y间叶片中SOD、POD、CAT、APX活性以及MDA含量在24h内基本呈现相反的变化趋势,而野牛草相连克隆分株O和Y间叶片中以上指标在24h内呈现趋于一致的变化规律。研究发现,野牛草酶促活性氧清除系统活性在一天内呈现节律性表达模式,且差异光周期处理下的野牛草相连克隆分株的活性氧清除系统的活性的节律性变化趋于同步。     

马尾松PmPGK1和PmGPIC基因的克隆和表达分析

为了解马尾松（Pinus massoniana）磷酸甘油酸激酶1（PGK1）与胞质溶胶葡萄糖磷酸异构酶（GPIC）的功能，采用RACE技术克隆了PmPGK1和PmGPIC基因，并进行了生物信息学分析与亚细胞定位，采用实时荧光定量PCR技术分析PmPGK1和PmGPIC的表达特性。结果表明，PmPGK1和PmGPIC全长为2 106和1 848 bp，分别编码507和566个氨基酸。PmPGK1和PmGPIC分别定位于叶绿体和胞质溶胶。PmPGK1表达量为新叶 > 老叶 > 新茎 > 根 > 花；而PmGPIC为老叶 > 花 > 新叶 > 新茎 > 根。低温胁迫24 h，PmPGK1和PmGPIC的表达量均随时间延长先降低后升高，且PmGPIC的表达量在处理2 h后即降至较低水平；高浓度CO₂胁迫24 h，PmPGK1的表达量随时间延长呈降低-升高-再降低的变化趋势，PmGPIC的表达下调但变化较不显著。因此，推测PmPGK1主要参与卡尔文循环及叶绿体/质体糖酵解，PmGPIC主要参与细胞质基质糖酵解；PmPGK1、PmGPIC活性在低温胁迫下均受抑制；PmPGK1活性在CO₂胁迫下受到显著抑制，而PmGPIC活性的影响不大。     

The seasonal cycle of Thermocyclops crassus (Fischer, 1853) (Copepoda: Cyclopoida) in a shallow,eutrophic lake

The seasonal cycle of Thermocyclops crassus was studied from 1985 to 1987, in the Gronne, a shallow, productive lake. T. crassus was present from late April to early October, while water temperature was above 10 °C. It produced three generations per year. Population peak was usually reached by the second generation, in August. Abundance was positively correlated with water temperature. Females carried 18.3 to 32.3 eggs on average. Reproduction rates were highest in July and August. Sex ratio was low, as females generally outnumbered males. Between October and April copepodites 4 went in diapause, predominantly in the deepest part of the lake and 0–4 cm deep in the bottom mud. Variations in body size were low across the year. T. crassus coexisted with the cyclopids, Cyclops vicinus and Mesocyclops leuckarti. In 1985 and 1986, abundances of M. leuckarti were low, while those of T. crassus and C. vicinus were high. In 1987 lower water temperatures, caused by cold weather, resulted in a marked decrease of the population density of T. crassus, while the abundance of M. leuckarti increased, and density of C. vicinus remained high. Comparison of the egg duration times of T. crassus, C. vicinus and M. leuckarti showed that at 15 °C and below T. crassus may be outcompeted by its comparatively longer egg development times. However, experiments in limnocorrals showed that T. crassus has an advantage over C. vicinus when fish predation is high.Department of Biology III, University of Ulm     

青藏高原两种植被类型净初级生产力与气候因素的关系及周转值分析

基于2008—2016年青海海北站9年净初级生产力及气候因子监测数据,分析了青藏高原高寒小嵩草草甸和高寒金露梅灌丛两种植被净初级生产力年际动态,并探讨了气候因子对其影响及其不同土层深度根系周转值特征。结果表明:(1)年际尺度上,小嵩草草甸地上净初级生产力表现为显著增加趋势,增幅为7.02 g m~(-2) a~(-1),而金露梅灌丛地上净初级生产力相对较为稳定;对于其地下净初级生产力和总生产力,小嵩草草甸和金露梅灌丛均表现为增加趋势(P0.05),9年间小嵩草草甸地上、地下和总净初级生产力平均值分别为(217.55±9.95)、(1882.75±161.33) g m~(-2) a~(-1)和(2100.30±163.38) g m~(-2) a~(-1),金露梅灌丛地上、地下和总净初级生产力9年间平均值分别为(256.27±11.4)、(1614.31±173.03) g m~(-2) a~(-1)和(1870.58±177.93) g m~(-2) a~(-1)。(2)不同植被类型地上净初级生产力对气候因素响应不同,金露梅灌丛地上净初级生产力主要受温度影响,而温度对小嵩草草甸地上净初级生产力无显著影响。此外,降水不是限制高寒生态系统草地地上净初级生产力主要因子,相比于降水影响,高寒生态系统地上净初级生产力更受温度调控。(3)年均温和年降水对金露梅灌丛和小嵩草草甸地下净初级生产力均无显著影响(P0.05),表明高寒生态系统,其地下生产力受外界气候条件变化影响微弱,是一个稳定的碳库。(4)两种植被类型其根系周转值均随着土壤深度的增加呈逐渐增加趋势,且高寒灌丛根系周转值明显高于高寒草甸根系周转值。研究表明,在全球气候变暖背景下将会增加金露梅灌丛地上净初级生产力,而对小嵩草草甸地上净初级生产力无显著影响。     

大兴安岭林区南、北部天然樟子松生长对气候变化的响应差异

利用树木年代学方法,建立大兴安岭林区南、北部樟子松(Pinus sylvestris var.mongolica)年轮宽度年表,探讨樟子松径向生长对气候变化的响应差异。结果表明,南部(阿尔山、海拉尔)树轮宽度主要与当年4—9月的平均标准化降水蒸散指数SPEI(Standardized Precipitation Evapotranspiration Index)极显著正相关(r=0.639,P0.01),而北部(漠河、塔河)树轮宽度主要与同时期的平均最低温极显著正相关(r=0.488,P0.01)。说明南部樟子松径向生长主要受当年4—9月的水分限制,北部主要受同期平均最低温调控。两个地区树木生长对降水的响应一致,对当年4—9月(6月除外)的温度响应相反。近几十年来随着温度显著升高(P0.01),南部树木生长对4—9月平均最高温的负响应不断增强,而北部树木对同时段平均最低温的正响应更加明显。同时,南部樟子松生长量快速下降(r=0.612,P0.001),而北部生长量显著增加(r=0.474,P0.001)。研究发现,高温加剧干旱胁迫是南部樟子松生长量下降的主要原因,而北部樟子松生长量增加是受到4—9月平均最低温和降水量的相互作用。如果持续变暖,未来樟子松分布区可能北移。