927 resultados para drought adaptability


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For the 6-year period 1987 through 1992, most of California suffered the worst or near-worst drought in a recorded history of about 140 years. Based on tree ring reconstructions, it may have been the worst in more than 400 years. The purpose of this paper is to review briefly the recent drought, then talk about the water supply situation this year, with some discussion of why the California drought is over hydrologically for most people; but for some, water supply problems continue.

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In 1993, enough rain and snow fell to end the 1987-1992 drought in California, at least in most areas. The 1994 season has again been dry and has given rise to questions about whether the drought has returned. But carryover reservoir storage from 1993 has been good and will be meeting a portion of the state's water needs this year. Concern about 1995 has placed California in a "drought watch" mode for 1994, a stage indicating caution but not yet a full drought.

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The introduction of the controlled hatchery system "Model CIFE D-81" resulted in a production of 60 lakhs Indian major carp seed in the drought area of eastern Uttar Pradesh. The hatchery system provided optimum temperature, dissolved oxygen and pH for breeding and hatching and also removed the metabolites generated. The technology is simple and the success of the program attracts pisciculturists.

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Photosynthetic activity during rehydration at four temperatures (5, 15, 25, 35 degrees C) was studied in a terrestrial, highly drought-tolerant cyanobacterium, Nostoc flagelliforme. At all the temperatures, the optimum quantum yield F-v/F-m increased rapidly within I It and then increased slowly during the process of rehydration. The increase in F-v/F-m at 25 and 35 degrees C was larger than that at 5 and 15 degrees C. In addition, the changes of initial intensity of fluorescence (F-0) and variable fluorescence (F-v) were more significant at 25 and 35 degrees C than those at 5 and 15 degrees C. Chlorophyll a content increased with the increase of temperature during the course of rehydration, with this being more pronounced at 25 and 35 degrees C. The photosynthetic rates at 25 and 35 degrees C were higher than those at 5 and 15 degrees C. Induction of chlorophyll fluorescence with sustained rewetting at 5 and 15 degrees C had two phases of transformation, whereas at 25 and 35 degrees C it had a third peak kinetic phase and showed typical chlorophyll fluorescence steps on rewetting for 24 h, representing a normal physiological state. A comparison of the chlorophyll fluorescence parameters, chlorophyll a content, and the chlorophyll fluorescence induction led to the conclusion that N. flagelliforme had a more rapid and complete recovery at 25 and 35 degrees C than that at 5 and 15 degrees C, although it could recover its photosynthetic activity at any of the four temperatures. (c) 2007 Published by Elsevier Ltd.

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利用400Gy 60Co γ射线处理87070谷子品系干种子,选育出辐谷7号新品种。该品种具有株型紧凑、茎秆矮、高产抗旱优质并且适应种植地区广等特点。2002年2月通过陕西省农作物品种审定委员会审定,截止2008年在黄土高原地区累计推广10.8万hm2。

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人类向大气中排放的大量氮氧化合物和氟氯烃类化合物(CFC’s)引起臭氧分子的分解,导致到达地球表面的紫外辐射增加,特别是UV-B辐射增强。本项目以青杨组杨树为模式植物,从形态和生理方面研究了来自不同UV-B背景下的康定杨与青杨在增强UV-B下的反应及其反应差异,并探讨了干旱、施肥对它们抗UV-B能力的影响。杨树具有分布广、适应性强、在生态环境治理和解决木材短缺方面均占有重要位置,研究成果可为生态系统的恢复与重建提供理论依据和科学指导。主要研究结果有以下: 1. 在温室中经过增强UV-B处理,杨树的外部形态及生理活动受到了一定程度的影响。增强UV-B导致康定杨、青杨的生物量、叶面积及节间长度降低,叶片增厚,SOD活性升高,膜伤害增加,而对叶片数目、R/S、叶绿素A、叶绿素B及整个叶绿素含量没有影响。两种杨树对UV-B胁迫的响应存在差异:在增强UV-B条件下,青杨的植株高度、生物量、叶面积、脯氨酸含量、长期用水效率受到的影响大于康定杨,相比而言,康定杨在比叶面积、叶片厚度、可溶性糖含量、UV-B吸收物质的含量及SOD和GPX活性方面增加的程度大于青杨。这些区别说明,来自于高海拔的康定杨比来自于低海拔的青杨对增强UV-B 具有更强的耐性。我们认为二者在叶片厚度、比叶面积、UV-B吸收物质含量及SOD、GPX活性差异是导致对增强UV-B耐性不同的原因。 2. 干旱与增强UV-B对杨树的生长和生理特性均产生了影响,而且两种胁迫共同作用时干旱表现减弱或加剧了UV-B对杨树某些形态和生理特性的影响。 据试验结果,干旱显著地降低了杨树的株高、叶片数目、叶面积,增加了叶片厚度,促进ABA的积累,提高了CAT活性。对于干旱,两种杨树之间也表现出了一定的差异性。可溶性蛋白质和脯氨酸在青杨叶片中得到显著积累,而在康定杨中没有变化。此外,CAT、长期用水效率在康定杨中受到的影响更加明显。长期用水效率的不同变化趋势说明两种杨树对水分胁迫采用了不同的用水策略,康定杨采用的是节水用水策略,提高用水效率,而青杨采用的是耗水的用水策略。根据干旱对叶面积、脯氨酸、ABA含量、CAT活性及长期用水效率等方面的影响,我们认为来自高海拔地区的康定杨比来自低海拔的青杨有更大的耐旱性,这是对生长环境长期适应的结果。在高海拔地区,因霜冻常带来土壤水分不可利用,降低了根系对水分的吸收,树木容易受到的生理性干旱。另外,高海拔的地区低的气温使植株对严寒有较强的耐性,减少了水分的需要。 生长于增强UV-B下的康定杨和青杨植株表现为高度降低,叶面积缩小,比叶面积增加;叶片栅栏组织、海绵组织均受到增强UV-B的影响,其厚度的增加导致整个叶片变厚。增强UV-B还显著提高了杨树的APX活性、UV-B吸收物质含量,而对叶片数目、ABA、可溶性蛋白质含量及CAT活性没有产生影响。试验中也观察到了两种杨树对增强UV-B响应的差异:与康定杨相比,在增强UV-B下青杨株高、叶面积降低的程度更大一些,SOD活性显著提高。另外UV-B吸收物质受到的影响不同。根据这些差别,高海拔的康定杨(3500 m)比来自低海拔的青杨(1500 m)增强UV-B有较强的耐性。 与水分充足情况下UV-B对植株的影响相比,干旱对杨树抗增强UV-B产生了一定的影响,表现为加剧或减弱UV-B对植物的影响,但这种影响与形态、生理指标有关。当干旱与增强UV-B共同作用时,杨树植株的株高、叶面积进一步降低、叶片进一步增厚。就脯氨酸的积累的而言,在没有水分胁迫时,增强UV-B促使它显著增加,而在干旱处理下这种效果变得不明显。干旱对增强UV-B的影响还与杨树的种类有一定的关系。在康定杨中,干旱减弱了增强UV-B对栅栏组织与海绵组织的影响,且在植株高度、叶面积上表现出累加效应,而在CAT上交互作用显著;但在青杨中干旱则加剧增强UV-B对栅栏组织与海绵组织的影响,在植株高度、叶面积及比叶面积上表现出显著的交互作用。据碳同素分析,在水分充足的条件下,无论是康定杨,还是青杨,增强UV-B均导致其长期用水效率的提高,然而当两种胁迫共同作用时,长期用水效率则表现出差异,在青杨中,长期用水效率得到进一步增高,而康定杨中干旱的效应被增强UV-B所减轻。 3. 田间试验表明,杨树的生长、生理特征都受到养分和增强UV-B的影响。施肥对杨树的影响表现为:提高了叶面积、生物量及SOD的活性,降低了抗坏血酸含量。对于施肥作用,两种杨树的反应也有区别:在康定杨中施肥显著增加了的叶片长度、宽度及光合色素的含量,降低了净光合速率、气孔导度及胞间CO2浓度;在青杨中,则SOD、GPX、APX活性表现增加。从试验看出,施肥对来自于高海拔地区的康定杨(3500 m)的影响较大,对来自低海拔的青杨(1500 m)影响较小,这与它们对原产地的生境适应有一定关系。在康定杨生长的高海拔地区,低温度和湿度不能为地上凋落物或土壤中的根分解提供理想的条件,造成当地土壤的低养分状况,所以当肥料施用以后,效果显著。 经过增强UV-B处理,杨树叶片中UV-B吸收物质含量、GPX的活性得到提高,而脯氨酸、丙二醛、可溶性蛋白质、叶绿素及类胡萝卜素含量没有受到影响。对于增强UV-B两种杨树受到的影响也有所不同:在青杨中增强UV-B导致叶面积缩小,生物量、净光合速率降低,APX的活性及长期用水效率的提高,而对康定杨的这些指标没有产生显著影响,相反抗氧化酶的活性明显高于青杨。这些差异性是由于两种杨树对原产地不同UV-B背景的长期适应结果。康定杨长期生长在较高UV-B环境中,对UV-B有较强的耐性。而青杨适应于较低的UV-B环境,对增强UV-B较为敏感。 试验中施肥也影响了植株对增强UV-B的反应,不过这种影响与杨树的种类及测定指标有一定的相关性。例如,在缺肥的情况下,青杨的长期用水效率和康定杨的叶绿素含量都受到增强UV-B的显著影响,而施肥以后这种影响变得不显著。在缺肥的条件下,GPX、APX在青杨中的活性、GPX在康定杨中的活性对增加UV-B反应不敏感;而施肥以后则变化显著,同样胞间CO2浓度在康定杨也有类似的变化。 For past decades, Ultraviolet radiation, especially UV-B reaching the Earth’s surface increased because of depletion of ozone layer resulted from emission of NxO and CFC’s from human activities. In this experiment, different species of Populus section Tacamahaca Spach from different UV-B background were selected as a model plant to assess the effects of enhanced UV-B radiation. Morphological and physiological traits induced by enhanced UV-B were observed and the different responses between P. kangdingensis and P. cathayana were discussed, furthermore the influences of drought and fertilizer on responses induced by enhanced UV-B were studied. Since poplars play an important role in lumber supply, and are important component of ecosystems due to their fast growth and wide adaptation, the study could provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem. The results are as follows: 1. The experiment conducted in a greenhouse indicated that morphological and physiological traits of two poplars were affected by enhanced UV-B radiation. Enhanced UV-B radiation not only reduced biomass, leave area and internode length, but also increased leaf thickness and SOD activity as well as MDA concentration and electrolyte rate. However, no significant changes in leaf numbers, root shoot ratio, and total chlorophyll and chlorophyll component were observed. There were different responses to enhanced UV-B radiation between two species. Compared with P. kangdingensis, cuttings of P. cathayana, exhibited lower height increment and smaller leaf area. In addition, there were significant differences in free proline, soluble protein, and UV-B absorbing compounds, and the activity of SOD and GPX, long-term WUE between them. Differences in plant height, biomass, leaf area, free proline concentration, and long-termed WUE showed that P. cathayana were more affected by enhanced UV-B radiation than P. kangdingensis. In contrast, more increase of specific leaf mass, leaf thickness, and soluble sugar, and UV-B absorbing compounds, and activity of SOD and GPX were observed in P. kangdingensis. According to these results, we suggested that P. kangdingensis from high elevation, which adapted to higher UV-B environments, had more tolerance to enhanced UV-B than P. cathayana from low elevation, which adapted to lower UV-B environment. We believe it was the difference of leaf thickness, specific leaf mass, and UV-B absorbing compounds as well as the activity of SOD and GPX resulted in lower adaptation of P. cathayana to enhanced UV-B radiation. 2. Growth and physiological traits of two poplars were affected by both drought and enhanced UV-B radiation. Moreover, it was observed that when two stresses applied together drought could exacerbate UV-B effects or decrease sensitivity to UV-B. In the experiment, drought significantly decreased plant height, leaf numbers, leaf area, and increased leaf thickness, and ABA, and CAT activity of two poplars. There were significant interspecific differences to drought stress. Exposed to drought, soluble protein and proline concentration were increased in P. cathayana but not in P. kangdingensis. However, more changes in CAT and long-term WUE were observed in kangdingensis. Different change in long-term WUE suggests that two poplars adapted different water-use strategies. P. kangdingensis employ a conservative water-use strategy, whereas P. cathayana employ a prodigal water-use strategy. Based on the differences in leaf area, accumulation of free proline and ABA, CAT activity as well as long-term WUE, we believed that P. kangdingensis from high elevation had a greater tolerance to drought than P. cathayana from low elevation,which is the result of adaptation to local environment. In high elevation area, trees are prone to suffer from physiological drought because of un-movable water caused by frost. Besides lower temperature enable the plants had greater adaptability to frost as a results the requirement of water is reduced Enhanced UV-B radiation decreased shoots height, leaf area, and increased specific leaf mass and thickness of palisade and sponge layer as well as APX activity and UV-B absorbing compounds in both species. Whereas, leaf numbers, ABA content, soluble protein and CAT activity showed no differences to enhanced UV-B radiation. Interspecific differences were also observed. Compared with P. kangdingensis, P. cathayana showed lower shoot height and smaller leaf area, higher SOD activity. Besides, variation in UV-B absorbing compounds was found. These differences suggested that P. kangdingensis from high elevation (3500 m) was more tolerant to enhanced UV-B radiation than P. cathayana from low elevation (1500 m). Compared with morphological and physiological changes induced by enhanced UV-B radiation under well-watered conditions, drought exacerbated or decreased these changes. However, these effects vary with parameters measured. When two stresses applied together, shoot height and leaf area further decreased while leaf thickness further increased. Under well-watered conditions, enhanced UV-B radiation significantly increased proline content, but such effect was not observed under drought conditions. The effect of drought on enhanced UV-B radiation was related to species. For example, drought reduced the effects of enhanced UV-B radiation on palisade parenchyma and sponge mesophyll in P. kangdingensis, and additive effects in shoot height and leaf area and interactive effect CAT activity were observed. In contrast, for P. cathayana drought significantly exacerbated the effects of enhanced UV-B radiation on palisade parenchyma and sponge mesophyll; there were noticeable interaction in shoot height, leaf area and specific leaf mass. As far as long-term WUE is concerned, it was increased by enhanced UV-B radiation under well-watered conditions in both species. While different effect was observed between two species in combination of two stresses. Long-term water use efficiency was further increased in P. cathayana whereas the effect was less significant in P. kangdingensis. 3. The field experiment showed that growth and physiological traits of poplars were affected by nutrition and enhanced UV-B radiation. Fertilization significantly increased leaf area, biomass and SOD activity, reduced Ascorbic acid concentration. There was interspecific difference in response to fertilization. For P. kangdingensis, fertilization significantly increased leaf width, leaf length and photosynthetic pigments content while net photosynthetic rate and stomatal conductance, intercellular CO2 concentration were significantly decreased. However, for P. cathayana, these parameters were unaffected except the increase of SOD, GPX and APX activity. From above, it could concluded that P. kangdingensis from high elevation was more affected by fertilization than P. cathayana, This difference was due to adaptation to local environment., The low temperature and moisture where P. kangdingensis was collected can not provided optimum to decompose roots and litter fall as a result the nutrition in soil was poor. Exposed to enhanced UV-B radiation, for both species UV-B absorbing compounds and GPX activity were significantly increased while proline, MDA, soluble protein, chlorophyll, carotenoids were not affected. Different responses were also observed between the two species. Enhanced UV-B radiation caused significant decreases in leaf area, biomass, net photosynthetic rate and increase in APX activity and long-term WUE in P. cathayana but not in P. kangdingensis. In addition, activity in antioxidant enzymes was much higher in P. kangdingensis than in P. cathayana. In the experiment fertilization affected responses of cuttings to enhanced UV-B radiation, but it concern species and parameters measured. Long-term WUE in P. cathayana and chlorophyll in P. kangdingensis were significantly increased by enhanced UV-B radiation under non-fertilization treatments while the increase was not found under fertilization treatment. In contrast, under no fertilization treatment enhanced UV-B radiation did not affected GPX and APX activity in P. cathayana and GPX in P. kangdingensis while significant increase appeared after application of fertilization. Similar effect of enhanced UV-B radiation on intercellular CO2 concentration in P. kangdingensis was observed.

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在青藏高原东部的亚高山针叶林区,如何尽快恢复这一生态脆弱地区的植被,改变生态环境恶化的趋势,是一个十分重要的课题。光一直被认为是植物种间相互替代,尤其是森林演替过程中植物相互替代或植被恢复中的关键环境要素之一。植物能否适应林冠下或林窗中异质的、或多变的光照条件,对其在林中的生存、分布、更新以及森林动态都是非常重要的。 本文以青藏高原东部亚高山针叶林的主要森林类型——岷江冷杉林群落的几种树苗为研究对象,采用实验生态学、生理及生物化学等方法,通过模拟针叶林不同大小林窗内光照强度的变化,在中国科学院茂县生态站内采用遮荫处理设置6个光照梯度(100、55、40、25、15与7%全光照),来研究具有不同喜光特性的植物对光强的响应与适应机制,其研究结果可为揭示亚高山针叶林的演替规律、以及人工林下幼苗的存活与定居提供科学依据,也能为苗木的生产与管理提供科学指导,尤其是对针阔树种在不同光强下的响应与适应的比较研究,能为如何将阔叶树种整合到人工针叶林中提供新的思路。 光强对植物生长的影响 光强对植物的生长具有重要作用,不同植物在各自适宜的光强梯度下才能生长良好。通过一个野外盆栽实验,来研究不同光强对植物生长的影响(第三章)。主要研究结果如下,低光强下植物株高/茎生物量增加,说明植物会将生物量更多用于高生长,以便有效地拦截光资源;在强光下,植物将生物量更多地向根部分配,使得植物在强光下能够吸收更多的水分,而避免干旱胁迫。 在第一个生长季节,以相对生长速率(RGR)表示,红桦和青榨槭在100%全光照下RGR最大,粗枝云杉在55%最大,岷江冷杉在25-40%下较好;然而,在第二个生长季节,2种阔叶树的相对生长速率(RGR)的适宜光强则变为25-55%,云杉为55-100%,而冷杉为25-100%。可见,从第一年到第二年,2种阔叶树苗更适宜在部分荫蔽的条件下生长;而2种针叶树苗对光的需求则逐渐增加,这可能是增加对根生物量相对投资的结果,因为以这种方式,强光下生长的针叶树幼苗更能保持其内部水分平衡,其生长不会因干旱胁迫而受到严重影响。另外,严重遮荫会引起冷杉幼苗死亡。 植物对光强的生理适应 植物可以通过自身形态和生理特征的调整,来发展不同的光能利用策略从而能够在林中共存。通过一个野外盆栽实验,研究了不同光强下生长的几种树苗的生理特征(第四章)对不同光强的响应与适应。结果显示:强光下,粗枝云杉和红桦的光合能力增加,而岷江冷杉和青榨槭在中度遮荫(25-55%)的条件下光合能力最大。植物叶氮和叶绿素含量增高,而光补偿点和暗呼吸速率降低,这些都是植物对低光环境的适应性反应;而强光下植物叶片和栅栏组织变厚,是对强光的一种保护性反应。 植物对光的可塑性反应 不同植物会表现出对光适应有利的生理和形态可塑性反应。本文对第三章、第四章的实验数据进行可塑性指数分析,来研究植物对光强的表型可塑性反应(第五章)。结果显示,生理特征调整是植物对不同光环境的主要适应途径。红桦和青榨槭的可塑性指数平均值要大于粗枝云杉和岷江冷杉,充分表明这2种阔叶树在生理和形态上较强的可塑性更有利于对光环境的适应,而具有比耐荫树种更强的适应能力。另外,2种针叶树相比,云杉的适应性更强。本研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说。 植物的光抑制与防御 当植物叶片吸收了过多光能,会发生光抑制现象。植物对光抑制的敏感性及防御能力对其生长具有重要意义。本文通过两个野外盆栽实验,研究了生长在强光下(第六章)和变化光强下(第八章)植物的光抑制现象及其防御策略。结果表明,在强光下或从遮荫状态转入强光下,植物都会发生光抑制,其对光抑制的敏感性与植物的耐荫性(或喜光)和演替状态有密切联系。长期生长在强光下的植物受到光抑制是可恢复的,而当处于荫蔽环境的植物突然暴露于强光下时,受到的光抑制不能完全恢复,可能是(部分)光合机构受到破坏的缘故。粗枝云杉和青榨槭防御光抑制伤害的能力较强,热耗散是其防御光抑制的主要途径。长期的强光作用能使岷江冷杉和红桦发生严重光抑制,甚至光伤害,而红桦能够通过“凋落老叶,萌发新叶”的途径来适应新的强光环境。 How to restore the vegetation of subalpine coniferous forest in eastern Qinghai-Tibet Plateau, and change the trend of ecological deterioration is a very important issue. Acclimation of tree seedlings to different and varing light environment affects to a great extent the successful regeneration and establishment of subalpine coniferous forests in southwestern China’s montane forest areas, because the ability to respond to such changing resource are commonly assumed to be critical to plant success, and have a growth advantage than others. In this paper, several species seedlings in Abies faxoniana community were chosed to study the response and adaptation to light intensity and the interspecific differences of adaptability in six shaded sheds (100, 55, 40, 25, 15 and 7% of full sunlight) in the Maoxian Ecological Station of Chinese Academy of Sciences. Our results could provide a strong theoretical evidence for understanding the forest succession laws of subalpine coniferous forests, and the survival and settlement of seedlings under plantations, and provide scientific direction for the production and management of seedlings, especially the comparative studies of the acclimation to light between the conifer and broadleaf trees could provide new ideas for how to integrate the broad-leaved trees into the artificial coniferous forest. Growth under different light intensity Light intensity plays an important role on plant growth. One field experiments was conducted to study the growth of tree seedlings of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii under different light intensities. The results showed that plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources, while biomass greater allocation to the roots, could make plants under high light environment absorb more water, and avoid drought stress. During the first growing season, the relative growth rates (RGRs) of Betula albo-sinensis and Acer davidii had the greatest values under the 100% of full light, for 55% of Picea asperata, and for 25-40% of Abies faxoniana. However, in the second growing season the the relative growth rates of the two broad-leaved trees changed and were appropriate for 25-55% of full light, for 55-100% of spruce, and for 25-100% of fir. Thus, from the first year to the second year, two broad-leaved seedlings maybe more suitable to partly shading environment, and two coniferous seedlings would have an increase in light demand, which may be an increased root biomass investment. Because in this way, seedlings grown under high light could better maintain their internal water balance, and thus its growth would not be seriously affected by drought stress. In addition, serious shading would cause fir seedlings to die. Acclimation of physiology to light Plants could coexist in forest ecosystem by forming different strategies of light use. One field experiments was conducted to study the acclimation of tree seedlings to different light intensity of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii. The results showed that the photosynthetic capacity of Picea asperata and Betula albo-sinensis exhibited a general tendency of increase with more light availability; but for Abies faxoniana and Acer davidii seedlings, their highest values of the same parameters were found under intermediate light regime (i.e. 25-55% of PFD relative to full sunlight). Plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources. Leaf nitrogen and chlorophyll content increased, while dark respiration rate and light compensation points decreased, all of which were adaptive response to the low light environment. On the contrary, plants under high light environment had the thicken leaves and palisade tissue, which was a protective response to high light. Phenotypic plasticity to light Phenotypic plasticity can be exhibited in morphological and physiological processes. Physiological characteristical adjustment is the main for plant adaptation to different light environment.The means of plasticity indexes for Betula albo-sinensis and Acer davidii seelings were greater than Picea asperata and Abies faxoniana, amplied that the two broad-leaved trees were much more adaptable to the environment. In addition, spruce had the higher adaptablity than fir. The findings supported the hypothesis that the ecological characteristics of the species determined the biological status and its biological habitat selection. Photoinhibition and photoprotection to light Compared with conifer, broad-leaved trees could better change leaf morphology and adjust biomass allocation to adapt to changing light environment. However, excess light can photoinhibit photosynthesis and may lead to photooxidative destruction of the photosynthetic appatus. Two field experiments were conducted to study the photoinhibition of photosynthesis. The results showed that when plants grown under high light environment or plants transferred from low to high irradiance, the four tree seedlings would undergo a period of photoinhibition. In four species, photoinhibited leaves could recover to initial photosynthetic rates when they were long-term planted under high light environment. However, when plants were suddenly exposed to high irradiance, this photoinhibition could not be reversible, may be the photosynthesis apparatus were (or partly) photooxidatively destructed.