辽东栎幼苗对干旱和去叶的生态反应的初步研究

幼苗阶段是植物种群生活史中亏损最严重的时期之一。通过干旱处理和去叶试验研究了辽东栎幼苗的生态反应。结果表明：⑴干旱处理显著降低了单叶面积、叶面积比和叶比重,从而也降低了单株生物量,但根比重却较对照组有显著上升;⑵去叶显著提高了单株叶数、叶面积比和叶比重,尽管单株生物量只稍高于对照组,这说明辽东栎在去叶后有一定的补偿能力。     

Interannual variation in precipitation and other planting conditions impacts seedling establishment in sown plant communities

Ecological restoration can reverse biodiversity loss worldwide, yet restoration goals and outcomes vary widely, which limits this potential. Divergent restoration outcomes may stem from variation in conditions at the outset of restoration, but empirical evidence is lacking and typically confounded with site differences. Additionally, precipitation is usually cited as the source of this variation, although a wide range of conditions can vary annually. We tested for effects of planting year on seedling establishment by installing identical restorations in three different years. Within those years, we manipulated rainfall with rain‐out shelters to disentangle the effects of precipitation from other annually variable conditions. Additionally, we tested whether increasing seed mix richness buffers against adverse planting conditions. For the first growing season after planting, we followed emergence and survival of sown prairie species and nonsown weed species to determine how planting year conditions influence an establishing plant community, if at all. We found that seedling establishment differed across planting years and precipitation treatments, and that varying emergence patterns by species led to differences in the composition of the first‐year community. We also found significant variation in sown species establishment across years when precipitation was held constant, illustrating the previously overlooked role of nonprecipitation drivers on planting year effects. Higher seed mix richness did not consistently improve establishment of sown species under different planting conditions. This research provides important experimental evidence for effects of interannual variation in planting conditions on first‐year establishment. Future work will examine how these initial changes affect longer‐term assembly dynamics.     

种子大小和干旱胁迫对辽东栎幼苗生长和生理特性的影响

在温室内遮阴条件下,设置80%、60%、40%和20%田间持水量(对照、轻度、中度和重度干旱)4个处理,研究种子大小和干旱胁迫对盆栽辽东栎幼苗生长和生理特性的影响。结果表明:大种子(3.05±0.38 g)幼苗的单株叶面积、总干质量和根冠比在所有处理均显著大于小种子(1.46±0.27 g)幼苗,前者的株高、基径、叶片数、比叶面积、相对生长率和净同化率等生长参数在轻度、中度和重度干旱处理均不同程度大于后者。大种子幼苗叶片过氧化物酶(POD)、过氧化氢酶(CAT)和超氧化物歧化酶(SOD)活性均高于小种子幼苗,前者叶片丙二醛(MDA)、可溶性蛋白、游离脯氨酸含量和叶绿素总量在部分干旱处理显著大于后者。除根冠比外,其他生长参数均随干旱胁迫增强逐渐减小,重度干旱处理大、小种子幼苗总干质量分别比对照降低19.4%和20.0%。POD、CAT和SOD活性均随干旱胁迫增强先升后降,在中度干旱处理,大、小种子幼苗POD活性分别显著高于对照126.7%和142.1%,CAT活性分别显著高于对照170.0%和151.9%。在重度干旱处理,大、小种子幼苗MDA含量分别显著高于对照86.5%和68.9%。可...     

不同密度下辽东栎幼苗子叶丢失及其对幼苗存活和生长的影响

在宁夏六盘山区龙潭林区华北落叶松(Larix principis-rupprechtii )人工林的不同样带, 按6个密度梯度(3.24、2.56、1.96、1.44、1.00和0.64株·m^-2)移栽萌发生根的辽东栎(Quercus wutaishanica)幼苗, 研究了啮齿动物对幼苗子叶取食的密度效应及其对幼苗存活与生长的影响。结果表明, 辽东栎幼苗的子叶留存率随着幼苗密度的增大而减小。2个高密度处理(3.24和2.56株·m^-2)的幼苗子叶留存率分别在移栽6周和5周后稳定在最低水平, 分别为8.64%和7.81%; 而2个较低密度处理(1.44和0.64株·m^-2)的幼苗在最后一次观测时子叶留存率仍很高, 分别为44.44%和31.25%, 且二者均显著高于其他密度处理(p < 0.05)。受啮齿动物取食的影响, 在从高到低的不同密度处理下, 幼苗的主根留存率分别为64.16%、80.46%、87.23%、80.25%、69.84%和69.84%; 在1.44株·m^-2处理下, 顶芽留存率最高(25.23%), 显著大于3.24株·m^-2处理(4.19%) (p < 0.05), 其他密度处理的顶芽留存率均不足20%。子叶被动物取食的幼苗和子叶完整幼苗的最终留存率基本一致, 仅在个别密度处理的某一观测时期差异显著; 顶芽被动物取食的幼苗留存率略低于顶芽完整幼苗的留存率, 但个别密度处理完全相反; 不同类型幼苗的留存率随时间推移而上下波动, 可能与幼苗顶芽被动物取食后萌生形成新芽有关。幼苗的株高、基径、叶片数和单株叶面积均随密度的减小而略有增大, 但除株高外, 其他3个生长参数均在1.44株·m^-2处理时最大, 与子叶是否被动物取食有一定的对应关系, 即高密度下更多的幼苗丢失了供应其生长所需营养物质的子叶, 从而影响了幼苗的早期生长。研究结果不仅丰富了“负密度效应”理论和种群更新的“增补限制”理论, 也可为六盘山区退化辽东栎灌丛和辽东栎次生林的恢复及生态系统管理提供 参考。     

The relative importance of carbohydrate and nitrogen for the resprouting ability of Quercus crispula seedlings

BACKGROUND AND AIMS: Plants need some kind of stored resources to resprout after shoot destruction. The aim of this study was to determine the relative importance of carbohydrate and nitrogen (N) storage levels for their ability to resprout. METHODS: A shoot clipping experiment was conducted on Quercus crispula seedlings, which were grown in a factorial experimental design, with two light levels (40% and 3% of full light) and three nutrient concentrations (low, medium and high). KEY RESULTS: At the time of shoot clipping (the end of spring leaf expansion), seedlings exposed to 40% light had an average total non-structural carbohydrate (TNC) concentration of 17.0% in their roots compared with 4.9% in the roots of seedlings exposed to 3% light, and the average amount of TNC (TNC pools) in the roots was 203.8 mg and 20.0 mg at 40% light and 3% light, respectively. In contrast, root N concentration averaged 2.3% in the 3% light treatment compared with 1.2% in the 40% light treatment, and it increased with successive rises in nutrient concentrations at both light levels. Regardless of the nutrient status, at the 40% light level >80% of the seedlings resprouted after shoot clipping. Few seedlings, however, resprouted at the 3% light level, particularly in the medium- and high-nutrient treatments. Furthermore, both root TNC concentrations and TNC pools decreased after resprouting, but the amount of root N remained constant. CONCLUSIONS: These results suggest that carbohydrate storage has a stronger influence on resprouting in Quercus crispula than N storage. However, the size of the resprouting shoot was positively correlated with the amount of both N and TNC in roots. The level of N storage is, therefore, also important for the growth of resprouting shoots.     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

Seed mix design and first year management influence multifunctionality and cost‐effectiveness in prairie reconstruction

Agricultural intensification continues to diminish many ecosystem services in the North American Corn Belt. Conservation programs may be able to combat these losses more efficiently by developing initiatives that attempt to balance multiple ecological benefits. In this study, we examine how seed mix design and first year management influence three ecosystem services commonly provided by tallgrass prairie reconstructions (erosion control, weed resistance, and pollinator resources). We established research plots with three seed mixes, with and without first year mowing. The grass‐dominated “Economy” mix had 21 species and a 3:1 grass‐to‐forb seeding ratio. The forb‐dominated “Pollinator” mix had 38 species and a 1:3 grass‐to‐forb seeding ratio. The grass:forb balanced “Diversity” mix, which was designed to resemble regional prairie remnants, had 71 species and a 1:1 grass‐to‐forb ratio. To assess ecosystem services, we measured native stem density, cover, inflorescence production, and floral richness from 2015 to 2018. The Economy mix had high native cover and stem density, but produced few inflorescences and had low floral richness. The Pollinator mix had high inflorescence production and floral richness, but also had high bare ground and weed cover. The Diversity mix had high inflorescence production and floral richness (comparable to the Pollinator mix) and high native cover and stem density (comparable to the Economy mix). First year mowing accelerated native plant establishment and inflorescence production, enhancing the provisioning of ecosystem services during the early stages of a reconstruction. Our results indicate that prairie reconstructions with thoughtfully designed seed mixes can effectively address multiple conservation challenges.     

Resprouting ability of<Emphasis Type="Italic"> Quercus crispula</Emphasis> seedlings depends on the vegetation cover of their microhabitats

To examine the effects of vegetation cover on the resprouting abilities of Quercus crispula seedlings, in each of three consecutive years, we artificially clipped seedlings growing in microhabitats with differing degrees of vegetation cover. We also investigated the relationship between the level of total nonstructural carbohydrate (TNC) and resprouting ability. Seedlings with clipped shoots in gaps produced larger resprouting shoots than those in the understory. Moreover, both the percentage of resprouting seedlings and the survival ratio in seedlings with clipped shoots were negatively correlated with the degree of vegetation cover. Seedlings stored high levels of TNC, especially in their roots, and their TNC levels were negatively correlated with the degree of vegetation cover. There were also positive relationships between the TNC levels in their roots and the degree of resprouting. Hence, we conclude that release from vegetation cover enhanced the resprouting ability of Q. crispula seedlings by increasing their levels of stored carbohydrate. The key variables affected were the resprouting ratio (the proportion of seedlings capable of producing new shoots) and the size of the resprouted shoots.     

生境变化对栓皮栎幼苗营养元素含量的影响

以栓皮栎在我国天然分布的南界、中部和北界为研究样点,将不同纬度的栓皮栎实生幼苗移栽到同一地点,于生长旺季同时在种源地和移栽地进行取样和测定,探讨生境变化对栓皮栎幼苗生长旺季营养元素含量特征的影响.结果表明:生境变化对栓皮栎幼苗各器官N含量和茎P含量的影响显著,而对各器官K含量和叶片、根P含量影响较小;在种源地,栓皮栎各器官的N含量与纬度间呈极显著的正相关,茎、根的全P含量随纬度升高而降低,叶片N/P的变化不明显;移栽后,北界栓皮栎幼苗各器官的N、P含量显著降低,不同纬度来源对栓皮栎幼苗N、P含量的影响更为明显;各纬度栓皮栎叶片的N/P均有不同程度的升高.不同纬度栓皮栎幼苗不同器官的营养元素含量存在差异,其对生境变化的响应也不相同.     

Top‐down control of rare species abundances by native ungulates in a grassland restoration

Ecological restorations are predicted to increase in species diversity over time until they reach reference levels. However, chronosequence studies in grasslands often show that diversity peaks after the first few years and then declines over time as grasses become more dominant. We addressed whether bison grazing and seed additions could prevent this decline in diversity. Exclosures that prevented bison grazing were compared with grazed plots over 4 years, and seed additions were conducted inside and outside exclosures to test for seed and microsite limitations. A previous study conducted 4‐months post seeding found that local species richness was primarily seed limited, but that grazing could sometimes increase seedling emergence. Here, we tested whether increased seedling emergence led to longer‐term increases in the species diversity of the plant community. We found that the seed addition effect grew smaller and the grazing effect grew stronger over time, and that seed additions affected the abundance of added species only when plots were grazed. Grazed plots had higher species diversity and lower biomass and litter buildup compared to non‐grazed plots. Our results suggest that moderate grazing by bison or management that mimics grazing can maintain diversity in grass‐dominated situations. Our results also emphasize the need to follow seed additions over several years to assess correctly whether seed limitation exists.     

黄土高原马栏林区辽东栎更新特性研究

研究了黄土高原马栏林区主要植被类型中的辽东栎幼苗的数量特征,更新层幼苗、幼树的实生和萌生特性及其辽东栎在垂直结构上的数量分布。结果表明:(1)辽东栎幼苗在黄土高原马栏林区分布广泛且数量充足。不同的植被类型中辽东栎种群表现出不同的大小级结构,在油松—辽东栎混交林、油松林和辽东栎林中辽东栎种群的幼苗、幼树和成树均占一定的比例,而在油松—白桦混交林、白桦林、山杨林和山杨—白桦混交林中辽东栎种群则以幼苗和幼树为主。表明辽东栎种群在该地区植被的发展过程中将产生重要的作用。(2)辽东栎在这一地区是由实生和萌生的个体混合组成的。在各种植被类型中实生植株的密度都高于萌生植株,辽东栎种群的更新在该地区可能主要是通过实生植株来完成的,即辽东栎实生植株在更新过程中起重要的作用。但萌生植株作为辽东栎顺利通过瓶颈的一种手段,作为辽东栎种群繁衍和稳定的一种途径,在辽东栎种群的更新中也起着一定的作用。     

土壤水分变化对长白山主要树种蒙古栎幼树生长的影响

选择长白山红松阔叶林主要优势树种蒙古栎为研究对象,人工控制3种施水量研究蒙古栎幼树形态、生物量效应和光合生理特征对土壤含水量变化的响应．结果表明,不同土壤含水量变化显著影响蒙古栎叶片、枝、根的生物量及其分配格局和叶片光合气体交换特征．水分胁迫改变幼树树冠结构,抑制幼树树高、地径、叶片大小、地上和地下生物量;同时,蒙古栎幼树根冠生物量比随着土壤水分含量的减少显著提高;供水量减少对幼树净光合速率、CO2利用率和碳利用率等特征有显著的负向影响;而叶片气孔导度、蒸腾速率和水分利用率对不同土壤含水量反应较复杂,只在土壤含水量较低时,幼树气孔导度、蒸腾速率明显降低,叶片水分利用率升高,表现出蒙古栎树种是干旱可变植物,长期水分胁迫可提高树种的耐旱能力．     

Evaluating outcomes of restoration ecology projects on limited budgets: assessment of variation in sampling intensity and sampling frequency for four habitat types

While best practices for evaluating restoration ecology projects are emerging rapidly, budget constraints often limit postrestoration monitoring, which emphasizes the need for practical and efficient monitoring strategies. We examined the postrestoration outcome for an ENGO (Nature Conservancy of Canada) project, to assess retroactively how variation in intensity and frequency of sampling would have affected estimates of plant species composition, diversity, and richness over time. The project restored four habitat types (mesic forest, oak woodland, wet meadow, and sand barren) using sculptured seeding of tallgrass prairie and woody species. Species‐level plant cover was monitored annually for 10 years in 168 2 × 2–m quadrats. We performed randomization tests to examine estimates of species diversity and richness as a function of the number of quadrats sampled, and assessed the necessity of annual sampling for describing changes in species composition and successional trajectories. The randomization tests revealed that sampling 10–17 quadrats, depending on habitat type, was sufficient to obtain estimates of species diversity that were at least 95% of values obtained from the whole dataset. Species richness as a function of number of quadrats sampled did not plateau, which suggests that rather than increasing the number of sampling quadrats, richness could be estimated more efficiently using nonquadrat based sampling techniques. Nonmetric multidimensional scaling analysis revealed that plant species composition largely stabilized by 3–5 years postrestoration depending on habitat type. By that time, native, seeded species dominated the restoration, and the benefits of annual sampling for tracking changes in species composition diminished.     

Effects of Frequent Mowing on Survival and Persistence of Forbs Seeded into a Species-Poor Grassland

Many early attempts at tallgrass prairie reconstruction failed to achieve the high species diversity of remnant prairies, and instead consist primarily of C₄ grasses. We hypothesized that frequent mowing of established prairie grasses could create sufficient gaps in the aboveground and belowground environment to allow for the establishment of native forbs from seed. We studied forb seedling establishment in a 25‐year‐old prairie planting in northern Iowa that was dominated by native warm‐season grasses. In winter 1999, 23 species of native forbs were broadcast into the recently burned sod at a rate of 350 viable seeds/m². Treatment plots were mowed weekly for either one or two growing seasons, and control plots were unmowed. Mowed plots had greater light availability than controls, especially when warm‐season grasses began to flower. Overwinter seedling mortality was 3% in mowed treatments compared to 29% in the controls. Forbs in mowed plots had significantly greater root and shoot mass than those in control plots in the first and second growing seasons but were not significantly more abundant. By the fourth growing season, however, forbs were twice as abundant in the mowed treatments. No lasting negative impacts of frequent mowing on the grass population were observed. Mowing a second year influenced species composition but did not change total seedling establishment. Experimental evidence is consistent with the idea that mowing reduced competition for light from large established grasses, allowing forb seedlings the opportunity to reach sufficient size to establish, survive, and flower in the second and subsequent years.     

不同分布区栓皮栎实生苗更新及其影响因子

分别在栓皮栎分布的边缘区黄土高原、半干旱核心区秦岭北坡以及湿润核心区秦岭南坡设置8块固定样地,调查了栓皮栎实生苗的年龄结构、生长状况、干质量累积与分配,并采用通径分析对影响实生苗更新的因子进行分析.结果表明:各分布区栓皮栎实生苗密度均随苗龄的增长而减小,且1～8年生实生苗密度均为秦岭南坡＞秦岭北坡＞黄土高原;各分布区相邻苗龄间的转化率差异显著,在黄土高原,7年生向8年生的转化率最低,为(30.2±2.9)％,在秦岭北坡和南坡,4年生向5年生的转化率最低,分别为(53.9±3.7)％和(50.0±2.1)％.随苗龄增长,各分布区实生苗的高度和干质量积累呈增加趋势,且秦岭南坡＞秦岭北坡＞黄土高原;主根长/株高逐渐下降,根幅/冠幅、根茎比则先升高后降低,三者均是黄土高原最高,秦岭南坡最低.温度、光照、林分郁闭度和灌层盖度是影响栓皮栎实生苗更新的直接因子,其中温度和光照起促进作用,郁闭度和灌层盖度起阻碍作用;而土壤速效氮含量和草本层盖度是重要的间接因子,分别起正向和负向作用.     

模拟淹水对池杉和栓皮栎光合生理生态过程的影响

池杉（Taxodium ascendens）和栓皮栎（Quercus variabilis）是我国亚热带河岸带和库塘消落区常见的植物,对于河岸带和库塘消落区的水土保持和水源涵养具有十分重要的作用。研究通过测定二者的最大净光合速率、光饱和曲线、二氧化碳饱和曲线、根／茎／叶的可溶性糖含量、叶片的比叶重（Ieaf mass per unit area,LMA）和根系活力等指标研究了模拟淹水（50d）对两年生栓皮栎和池杉树苗生理生态过程的影响。在淹水过程中,栓皮栎的最大净光合速率不断下降,到试验结束时淹水处理的最大净光合速率降低为对照的9％,而且处理与对照差异极显著;而在整个淹水过程中池杉则在处理和对照间没有显著差异。淹水胁迫下：（1）淹水导致栓皮栎表观量子效率（φ）不断下降,淹水第6天降为对照的59．9％左右（P〈0．001）,第26天降为对照的40．9％左右（P〈0．001）,而26d的淹水对池杉的光饱和曲线却影响不大;（2）淹水导致栓皮栎羧化效率（CE）的不断下降,在正常生长条件下栓皮栎的CE为（0．331&#177;0．075）μmolm^-2s^-1Pa^-1。淹水第10天栓皮栎的CE下降到对照的22．1％,淹水第30天降到对照的16．8％,且差异均显著（P〈0．001）。淹水对池杉的二氧化碳饱和曲线没有造成显著影响;（3）淹水导致两个种的根／茎／叶可溶性糖含量呈现动态变化。50d的淹水导致栓皮栎叶片可溶性糖含量相比于对照略有上升,茎和根可溶性糖含量显著上升。而池杉根／茎／叶可溶性糖含量对照和处理之间差异不显著;（4）淹水导致栓皮栎的比叶重持续升高,池杉的与对照没有显著差异;（5）淹水导致栓皮栎的根系活力持续下降,池杉则开始时下降后又升高,这可能与池杉在淹水过程中产生了有利于吸收氧气的不定根,而栓皮栎没有产生不定根有关。以上实验结果说明相对池杉,栓皮栎应属于对淹水较敏感的树种。短期淹水（1～10d）导致栓皮栎光合速率下降的生理生态原因应该至少包括表观量子效率和羧化效率下降两个原因。而长期淹水条件下,叶片碳水化合物的积累（LMA升高）导致的负反馈效应也可能是光合速率下降的原因之一。     

模拟酸雨胁迫对青冈幼苗光合特性和叶绿素荧光参数的影响

选择亚热带常绿阔叶林优势树种青冈幼苗为研究对象,设置3个酸雨处理：pH 2.5、pH 4.0、pH 5.6(CK),研究不同强度的模拟酸雨对青冈幼苗光合特性、叶绿素荧光参数和叶绿素含量的影响.结果表明：经过2年酸雨处理,青冈的净光合速率随着酸雨强度的增加而显著上升.pH 2.5、pH 4.0处理的酸雨增大了青冈的气孔导度和蒸腾速率,且对pH 2.5处理的影响更为显著.胞间CO₂浓度的大小顺序是pH 2.5>pH 5.6>pH 4.0.pH 2.5、pH 4.0处理青冈的最大净光合速率、光补偿点、光饱和点、暗呼吸速率显著高于对照,表观量子效率对酸雨胁迫不敏感.pH 2.5、pH 4.0处理青冈的PSⅡ原初光能转化效率和PSⅡ的潜在活性显著高于对照.青冈的叶绿素相对含量大小顺序是pH 2.5>pH 5.6>pH 4.0,且pH 2.5与pH 4.0之间有显著差异.说明青冈幼苗的光合参数、叶绿素荧光参数指标在pH 2.5和pH 4.0的酸雨处理下有所增加,且在pH 2.5强度下增加更为明显.     

辽东栎(Quercus liaotungensis)幼苗对土壤干旱的生理生态适应性研究

用盆栽方法人工模拟土壤干旱条件,研究辽东栎天然萌生幼苗对土壤干旱的生理生态反应。结果表明:随土壤含水量的减少,辽东栎幼苗的耗水量明显下降,耗水高峰期提前,在重度干旱下耗水峰形由单峰变为双峰。在干旱胁迫前期辽东栎叶水势变化较平缓,后期则急剧下降,尤其在严重干旱下,水势变化趋势呈"M"形,叶片含水量较稳定,对土壤水分含量变化不敏感。在中度干旱下辽东栎叶片的持水力有所增加。辽东栎属低蒸腾速率树种,平均仅为2.98μg cm^-2·s^-1。不同生长季节蒸腾速率日进程不同, 8月份的蒸腾速率日进程在适宜水分下是双峰曲线,在中度干旱下是单峰曲线,重度干旱下蒸腾速率一直维持在较低水平,呈波状起伏;9月份的蒸腾速率下降近50%,仍有明显的单峰(适宜水分下)和双峰(中度干旱下)。对蒸腾速率与环境因子的简单相关分析表明:在适宜水分和中度干旱下,光照强度对辽东栎幼苗蒸腾速率影响最大,在重度干旱下,大气温度对辽东栎的蒸腾速率影响较大。随土壤含水量减少,辽东栎幼苗的蒸腾速率在中度干旱时上升,重度干旱时急剧下降,光合速率、瞬时水分利用效率、羧化效率均下降;地上部生长受抑,根冠比加大, WUE₁下降,而WUE₂在中度干旱下升高,在重度干旱下下降。其幼苗不耐高温强光,高温强光对其光合和蒸腾有抑制作用,特别是在土壤严重干旱下表现更明显。