不同生长阶段沙地彰武松与樟子松抗逆生理特性比较研究
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摘要
为客观评价彰武松(Pinus densiflora var.zhangwuensis)的抗逆性及更好的理解其抗性机理,本次研究以4a生彰武松和樟子松(P. sylvestris var.mongolica)为试材开展了抗逆性试验。通过盆栽控水试验,对彰武松、樟子松苗木在不同水分梯度下形态指标和叶片内丙二醛(MDA)、过氧化氢酶(CAT)、叶绿素(Chl)、脯氨酸(Pro)和叶片含水率(LWC)5项抗旱生理指标进行了测定。结果表明:彰武松总干重、根干重、茎干重和根茎比等四个指标都显著高于樟子松,而单片针叶面积显著小于樟子松,针叶厚度略大。在不同水分梯度下,彰武松的MDA含量都小于樟子松,CAT活性、Pro含量都高于樟子松。轻度干旱区(40%FMC, SWP=-4.82bar)樟子松叶片中MDA、CAT和Pro较对照区(100%FMC, SWP=-2.81bar)均呈现上升趋势,在中度干旱区(30%FMC,
SWP=-10.35bar)这三个指标继续明显升高,在重度干旱区(20%FMC, SWP=-20.41bar)它们达到了最大值;而轻度干旱区中彰武松叶片的这三个生理指标反而低于对照区,在中度干旱区它们才开始增大,尤其是CAT活性较对照区显著增加,这些指标也在重度干旱区中达到了最大值。这表明彰武松幼苗在更低的土壤含水率下才表现出胁迫伤害,并且在开始受到胁迫伤害时就迅速提高了保护酶(CAT)活性,增加了渗透调节物质含量(Pro),实现了对干旱胁迫较强的忍耐性和较好的适应性。在不同水分梯度下,彰武松Chl含量和LWC均大于樟子松,这两个指标在同一树种中的变化趋势也相同,大小排序均为:对照区>轻度干旱区>中度干旱区>重度干旱区。
本次研究通过采用盆栽试验法研究NaCl、Na2CO3、NaHCO33种单盐胁迫及NaOH碱胁迫对苗木生长和叶片生理生化指标的影响。栽培基质为章古台当地细沙,在设置盐胁迫处理时,使基质总盐量为4g?kg-1;设置NaOH胁迫,使基质pH值达9.0。结果表明:彰武松苗木在盐碱胁迫下受害级别较小,根系忍耐指数(RTI)较大。盐碱胁迫下彰武松叶片丙二醛(MDA)含量没有显著提高,而在胁迫诱导下,过氧化氢酶(CAT)活性显著提高,最大增幅是对照的22.6倍。盐碱胁迫下彰武松叶片叶绿素(Chl)含量降幅较小,叶片含水率(LWC)有所提高。樟子松与彰武松表现出了不同规律,盐碱胁迫下其MDA含量均提高(NaCl处理)和显著提高(其他三个处理),但其CAT活性没有显著提高。盐碱胁迫下樟子松叶片Chl含量显著降低,Na2CO3、NaHCO3及NaCl处理下樟子松Chl含量分别是对照的36.2%、21.7%和62.3%。另外,胁迫下樟子松LWC有所下降。初步结论:彰武松通过提高CAT活性和维持较高Chl和LWC含量保证了其较高的耐盐碱性,其耐盐碱性高于樟子松。彰武松针叶内Fe元素含量较高是其保持较高CAT活性和Chl含量的一个原因,而Zn、Cu元素含量较高也与其抗性强有关。对于彰武松来说,盐碱胁迫并未造成干旱胁迫,因此其生长受到的抑制并不是干旱胁迫造成的,而是由盐碱胁迫引发的离子毒害和养分不平衡等原因造成的。
本次研究通过采用冰箱冷冻方法,对两个树种开展了不同温度梯度的抗寒胁迫试验。将电导法和生理生化法相结合,通过对胁迫前后苗木叶片相对电导率和生理生化指标(MDA、CAT、Pro、可溶性糖SS)进行测定,试图鉴定两个树种的抗寒性并明确它们的抗寒机理。结果表明:经抗寒性锻炼后,彰武松和樟子松的半致死温度分别为-54.23℃和-50.34℃,表明二者均是抗寒性较强的树种,低温适应后彰武松抗寒性略高于樟子松。伴随胁迫程度的加深,两个树种的Pro和SS变化模式基本相同,而CAT活性变化模式有所不同。在低于-20℃之后,彰武松CAT活性逐渐升高,而樟子松则保持恒定。在各温度梯度下,彰武松Pro和SS含量低于樟子松,而CAT活性高于樟子松。彰武松和樟子松抗寒机理略有不同:从各指标数量上看,彰武松侧重于启动抗氧化酶活性而抵御胁迫伤害,而樟子松侧重于利用渗透调节能力来增强抗寒性。从各指标变化随胁迫程度的变化趋势看,在轻度低温胁迫下,两个树种均通过提高渗透调节物质(Pro和SS)含量来增加抗寒性;在能造成胁迫伤害的重度低温胁迫下(-40℃~-60℃),彰武松提高了CAT活性,并且维持了SS增量,使其MDA含量增幅较小,而樟子松没有这样的机制。
本次研究还采用Li-6400光合测定系统对性成熟(18a)阶段彰武松和樟子松光合及蒸腾指标不同季节日变化进行了测定,并采用切枝蒸腾法对两个树种叶片气孔蒸腾和角质层蒸腾进行对比测定,评价了气孔开闭敏感性,探讨了两个树种光合生产与蒸腾耗水特性。结果表明:在同样生境条件下,彰武松比樟子松有较大的光合速率(Pn)和较小的蒸腾速率(Tr)。在5月和7月,彰武松的Pn和Tr日变化呈现明显双峰型,其Pn和Tr“午休”现象均主要受气孔限制;在10月呈单峰型。樟子松的Pn和Tr日变化在整个生长季均呈单峰型,而且,彰武松日光合量(DAP)均高于樟子松,是樟子松的163.4%(5月)、211.1%(7月)和183.6%(10月)。光响应曲线参数表明:在不同月份,彰武松最大光合速率(Pmax)均大于樟子松,且光饱和点(LSP)较高,光补偿点(LCP)较低。在任意被测时刻,彰武松气孔导度(Gs)和Tr都小于樟子松。彰武松具有较小气孔和角质层蒸腾速度,并且在同样干旱条件下,彰武松气孔下陷,其气孔的开闭反应更加敏感。彰武松水分利用效率(WUE)较高,约是樟子松的2.29倍。彰武松单位面积叶氮含量(Narea)和光合氮素利用效率(PNUE)较高,且叶肉胞间气相导度(gias)较大。彰武松气孔直径和密度均较小也是其能保持较小Gs、Tr,从而保持较大WUE的一个原因。这些结果暗示,彰武松以其高的光合速率和低的蒸腾耗水特性,提高水分利用效率,以其敏感的气孔开闭机制和旱生叶片结构进而实现在干旱半干旱地区的速生特性。
For an objective evaluation of stress resistance and a better understanding of its mechanisms in Pinus densiflora var. zhangwuensis, four-year-old P. densiflora var. zhangwuensis and P. sylvestris var. mongolica was adopted as test materials, and experimental researches on adversity resistance were carried out. With a potting experiment, morphological parameters and five physiological indicators including malondialdehyde(MDA), catalase(CAT), chlorophyll(Chl), proline(Pro), and leaf water content(LWC) of P. densiflora var. zhangwuensis and P. sylvestris var. mongolica were measured in different soil moisture gradients. The results showed that: P. densiflora var. zhangwuensis had significantly heavier total dry weight, dry weight of root, dry weight of stem and greater ratio of root and stem, had significantly smaller single needle area, and had slightly more thick needle than P. sylvestris var. mongolica. In different water gradients, P. densiflora var. zhangwuensis had lower MDA contents, and higher CAT and Pro contents than P. sylvestris var. mongolica. In mild drought area(40%FMC, SWP=-4.82bar), MDA, CAT and Pro contents of P. sylvestris var. mongolica had a ascending trend in comparison with those in control area(100%FMC, SWP=-2.81bar), and the three physiological indicators had a further increase in moderate drought area(30%FMC, SWP=-10.35bar) and reached to the maximum in serious drought area(20%FMC, SWP=-20.14bar). By contrast in mild drought area these three physiological indicators in leaves of P. densiflora var. zhangwuensis were lower than those in control area, and began to increase in moderate drought area, especially the CAT content was significantly higher than that in control area, and they also reached to the maximum in serious drought area. The above results determinated that P. densiflora var. zhangwuensis showed stress symptoms at a lower soil moisture, and it could endure and adapt to drought stress by improving protective enzyme activity(CAT) and increasing osmotic substance(Pro). At different moisture gradients, both Chl and LWC of P. densiflora var. zhangwuensis were more than those of P. sylvestris var. mongolica, and these two indicators in the same species had the same changing trend,which was : CK > mild drought area > moderate drought area > serious drought area.
For an objective evaluation of salinity-alkalinity tolerance and a better understanding of its mechanisms in Pinus densiflora var. zhangwuensis, 4-year-old P. densiflora var. zhangwuensis and P. sylvestris var. mongolica were studied. Pot experiments were used to examine the effects of three types of salt (NaCl, Na2CO3, and NaHCO3) and alkali stress (NaOH) on seedling growth and physiological and biochemical indices. The substrate was fine sand taken from Zhanggutai sandy land, Liaoning Province, China. In the salt stress treatments, the matrix salt content was 4 g salt kg-1, and in NaOH stress treatments, the matrix pH value was 9.0. In P. densiflora var. zhangwuensis, the injury level was smaller, and the root tolerance index was bigger under salt-alkali stress. Malondialdehyde (MDA) content did not significantly increase under salt-alkali stress, while catalase (CAT) activity was significantly induced by stress, by as much as 22.6 times that of the control. Finally, chlorophyll (Chl) content did not significantly decline, and leaf water content (LWC) increased slightly. Pinus sylvestris var. mongolica responded differently to salt-alkali stress; its MDA content increased (NaCl treatment) or significantly increased (remaining three treatments), but its CAT activity did not increase significantly. Leaf Chl content declined significantly under salt-alkali stress, with Chl contents in the Na2CO3, NaHCO3, NaCl treatments 36.2%, 21.7%, and 62.3% lower than in the control. In addition, LWC decreased in leaves under stress. The results suggest that P. densiflora var. zhangwuensis has greater salt-alkali tolerance than P. sylvestris var. mongolica and that it ensures this tolerance by increasing CAT activity and maintaining high contents of Chl and leaf water. Higher levels of iron in needles of P. densiflora var. zhangwuensis enhanced CAT activity and Chl content, while higher levels of elemental zinc and copper were also associated with stronger resistance. In P. densiflora var. zhangwuensis, salinity stress did not cause drought stress, so growth was not inhibited by drought stress, but by ion toxicity and nutrient imbalance due to salinity stress.
Freezer method was used to carry out cold-stress test at different temperature gradients. With the combination between conductivity method and physiological method, relative conductivity as well as physiological and biochemical indicators such as MDA, CAT, Pro and SS in leaves of seedlings before and after stress were measured in order to identify cold resistance of the two species and to clarify their mechanism of cold hardiness. The results showed that: After cold resistance exercise, half-lethal temperature of P. densiflora var. zhangwuensis and P. sylvestris var. mongolica was -54.23℃and -50.34℃, respectively, which indicated that both species were strong cold hardiness, but a little cold hardiness in P. densiflora var. zhangwuensis higher than P. sylvestris var. mongolica due to cold acclimation. With the deepening of stress, Pro and SS changes model of two species was basically the same, while CAT activity in different patterns. In lower than -20℃, the CAT activity in leaves of P. densiflora var. zhangwuensis seedlings gradually increased, while that of P. sylvestris var. mongolica remained constant. In every temperature gradient, Pro and SS content was lower in leaves of P. densiflora var. zhangwuensis than P. sylvestris var. mongolica, while CAT activity of the former higher than the latter. The two species had slightly different cold-resistance mechanisms: From the point of view of content of indicators, P. densiflora var. zhangwuensis focused on using higher antioxidant enzyme activities to resist stress injury, and P. sylvestris var. mongolica focused on using higher ability of osmotic adjustment to enhance cold tolerance. From the point of view of variation trend in the indicators with the stress deepening, in mild cold stress, the two species could increase cold hardiness by increasing osmolyte (Pro and SS) content. But in severe cold stress (-40℃~-60℃) that caused stress injuries, P. densiflora var. zhangwuensis had a smaller increase of MDA by increasing CAT activity and maintaining the SS increments, while P. sylvestris var. mongolica had no such mechanism.
In order to photosynthesize, land plants must open their stomata to exchange small amounts of CO2 at the cost of losing a lot of water vapor through transpiration. This gas exchange is unequal in that more water is lost than CO2 is taken in. In general, fast growing species are characterized by their high photosynthetic capacity, most of which is achieved at the expense of consuming large amounts of water. This trade-off poses a serious challenge to forestry in the selection of fast-growing tree species in arid and semiarid areas. The ideal forestry species for drier climates should maintain a high photosynthetic capacity as well as low water consumption for transpiration, but such species are very rare. In this study, the LI-6400 (LICOR, Inc. Lincoln, NE, USA) portable photosynthesis system was employed to measure diurnal changes in photosynthetic and transpiration indices in sexually mature P. densiflora var. zhangwuensis and P. sylvestris var. mongolica (18 years old). In addition, the cut-branch transpiration method was used to compare stomatal and cuticular transpiration in order to evaluate the sensitivity of stomatal opening and closing and to characterize the photosynthetic productivity and water consumption for transpiration in these two species in different growing seasons. P. densiflora var. zhangwuensis had a higher photosynthesis rate (Pn) and a lower transpiration rate (Tr) than P. sylvestris var. mongolica under the same conditions. In May and July, the diurnal changes of Pn and Tr in P. densiflora var. zhangwuensis formed double-peaked curve as a result of mid-day stomatal limitation, while it formed single-peaked curve in October. The daily variation of Pn and Tr in P. sylvestris var. mongolica exhibited a monopeak curve throughout the growing season. The net photosynthesis per day in P. densiflora var. zhangwuensis was higher than in P. sylvestris var. mongolica; values for the former were 163.4 (May), 211.1 (July), and 183.6 (October) percent of the latter. The photoresponse parameter measurements showed that, in different months, the maximum rate of photosynthesis of P. densiflora var. zhangwuensis was greater than that of P. sylvestris var. mongolica, and the former also had a higher light saturation point and a lower light compensation point. The stomatal conductivity (Gs) and Tr of P. densiflora var. zhangwuensis were lower than those of P. sylvestris var. mongolica when measured at random times. The stomatal and cuticular Tr of P. densiflora var. zhangwuensis were lower, and in the same drought conditions, its stomata were deeper and had a higher sensitivity for opening and closing. Water use efficiency (WUE) of P. densiflora var. zhangwuensis was 2.29 times that of P. sylvestris var. mongolica. Analysis of the correlation between Gs and WUE showed that P. densiflora var. zhangwuensis could maintain high WUE when Gs levels were high. In May and July, when P. densiflora var. zhangwuensis exhibited midday photosynthetic depression, its Gs level was low, usually in the 90~200 mmol ? m-2 ? s-1 range. At the same time, its WUE was higher than that of P. sylvestris var. mongolica, indicating that P. densiflora var. zhangwuensis could effectively conserve water by closing its stomata rapidly at midday, so as to maintain its high WUE. Nitrogen content per unit leaf area(Narea) and photosynthetic nitrogen use efficiency (PNUE) was higher, and gaseous conductance through intercellular air space(gias) was also larger in P. densiflora var. zhangwuensis. The stomata diameter and density was small, which was also able to maintain a smaller Gs and Tr, and thus a reason to maintain a larger WUE in P. densiflora var. zhangwuensis. These results implied that P. densiflora var. zhangwuensis can improve its WUE, yielding its higher Pn and lower Tr, and achieves its fast growth in arid and semiarid regions through its sensitive stomatal response and leaf xeromorphism.
SWP=-10.35bar)这三个指标继续明显升高,在重度干旱区(20%FMC, SWP=-20.41bar)它们达到了最大值;而轻度干旱区中彰武松叶片的这三个生理指标反而低于对照区,在中度干旱区它们才开始增大,尤其是CAT活性较对照区显著增加,这些指标也在重度干旱区中达到了最大值。这表明彰武松幼苗在更低的土壤含水率下才表现出胁迫伤害,并且在开始受到胁迫伤害时就迅速提高了保护酶(CAT)活性,增加了渗透调节物质含量(Pro),实现了对干旱胁迫较强的忍耐性和较好的适应性。在不同水分梯度下,彰武松Chl含量和LWC均大于樟子松,这两个指标在同一树种中的变化趋势也相同,大小排序均为:对照区>轻度干旱区>中度干旱区>重度干旱区。
本次研究通过采用盆栽试验法研究NaCl、Na2CO3、NaHCO33种单盐胁迫及NaOH碱胁迫对苗木生长和叶片生理生化指标的影响。栽培基质为章古台当地细沙,在设置盐胁迫处理时,使基质总盐量为4g?kg-1;设置NaOH胁迫,使基质pH值达9.0。结果表明:彰武松苗木在盐碱胁迫下受害级别较小,根系忍耐指数(RTI)较大。盐碱胁迫下彰武松叶片丙二醛(MDA)含量没有显著提高,而在胁迫诱导下,过氧化氢酶(CAT)活性显著提高,最大增幅是对照的22.6倍。盐碱胁迫下彰武松叶片叶绿素(Chl)含量降幅较小,叶片含水率(LWC)有所提高。樟子松与彰武松表现出了不同规律,盐碱胁迫下其MDA含量均提高(NaCl处理)和显著提高(其他三个处理),但其CAT活性没有显著提高。盐碱胁迫下樟子松叶片Chl含量显著降低,Na2CO3、NaHCO3及NaCl处理下樟子松Chl含量分别是对照的36.2%、21.7%和62.3%。另外,胁迫下樟子松LWC有所下降。初步结论:彰武松通过提高CAT活性和维持较高Chl和LWC含量保证了其较高的耐盐碱性,其耐盐碱性高于樟子松。彰武松针叶内Fe元素含量较高是其保持较高CAT活性和Chl含量的一个原因,而Zn、Cu元素含量较高也与其抗性强有关。对于彰武松来说,盐碱胁迫并未造成干旱胁迫,因此其生长受到的抑制并不是干旱胁迫造成的,而是由盐碱胁迫引发的离子毒害和养分不平衡等原因造成的。
本次研究通过采用冰箱冷冻方法,对两个树种开展了不同温度梯度的抗寒胁迫试验。将电导法和生理生化法相结合,通过对胁迫前后苗木叶片相对电导率和生理生化指标(MDA、CAT、Pro、可溶性糖SS)进行测定,试图鉴定两个树种的抗寒性并明确它们的抗寒机理。结果表明:经抗寒性锻炼后,彰武松和樟子松的半致死温度分别为-54.23℃和-50.34℃,表明二者均是抗寒性较强的树种,低温适应后彰武松抗寒性略高于樟子松。伴随胁迫程度的加深,两个树种的Pro和SS变化模式基本相同,而CAT活性变化模式有所不同。在低于-20℃之后,彰武松CAT活性逐渐升高,而樟子松则保持恒定。在各温度梯度下,彰武松Pro和SS含量低于樟子松,而CAT活性高于樟子松。彰武松和樟子松抗寒机理略有不同:从各指标数量上看,彰武松侧重于启动抗氧化酶活性而抵御胁迫伤害,而樟子松侧重于利用渗透调节能力来增强抗寒性。从各指标变化随胁迫程度的变化趋势看,在轻度低温胁迫下,两个树种均通过提高渗透调节物质(Pro和SS)含量来增加抗寒性;在能造成胁迫伤害的重度低温胁迫下(-40℃~-60℃),彰武松提高了CAT活性,并且维持了SS增量,使其MDA含量增幅较小,而樟子松没有这样的机制。
本次研究还采用Li-6400光合测定系统对性成熟(18a)阶段彰武松和樟子松光合及蒸腾指标不同季节日变化进行了测定,并采用切枝蒸腾法对两个树种叶片气孔蒸腾和角质层蒸腾进行对比测定,评价了气孔开闭敏感性,探讨了两个树种光合生产与蒸腾耗水特性。结果表明:在同样生境条件下,彰武松比樟子松有较大的光合速率(Pn)和较小的蒸腾速率(Tr)。在5月和7月,彰武松的Pn和Tr日变化呈现明显双峰型,其Pn和Tr“午休”现象均主要受气孔限制;在10月呈单峰型。樟子松的Pn和Tr日变化在整个生长季均呈单峰型,而且,彰武松日光合量(DAP)均高于樟子松,是樟子松的163.4%(5月)、211.1%(7月)和183.6%(10月)。光响应曲线参数表明:在不同月份,彰武松最大光合速率(Pmax)均大于樟子松,且光饱和点(LSP)较高,光补偿点(LCP)较低。在任意被测时刻,彰武松气孔导度(Gs)和Tr都小于樟子松。彰武松具有较小气孔和角质层蒸腾速度,并且在同样干旱条件下,彰武松气孔下陷,其气孔的开闭反应更加敏感。彰武松水分利用效率(WUE)较高,约是樟子松的2.29倍。彰武松单位面积叶氮含量(Narea)和光合氮素利用效率(PNUE)较高,且叶肉胞间气相导度(gias)较大。彰武松气孔直径和密度均较小也是其能保持较小Gs、Tr,从而保持较大WUE的一个原因。这些结果暗示,彰武松以其高的光合速率和低的蒸腾耗水特性,提高水分利用效率,以其敏感的气孔开闭机制和旱生叶片结构进而实现在干旱半干旱地区的速生特性。
For an objective evaluation of stress resistance and a better understanding of its mechanisms in Pinus densiflora var. zhangwuensis, four-year-old P. densiflora var. zhangwuensis and P. sylvestris var. mongolica was adopted as test materials, and experimental researches on adversity resistance were carried out. With a potting experiment, morphological parameters and five physiological indicators including malondialdehyde(MDA), catalase(CAT), chlorophyll(Chl), proline(Pro), and leaf water content(LWC) of P. densiflora var. zhangwuensis and P. sylvestris var. mongolica were measured in different soil moisture gradients. The results showed that: P. densiflora var. zhangwuensis had significantly heavier total dry weight, dry weight of root, dry weight of stem and greater ratio of root and stem, had significantly smaller single needle area, and had slightly more thick needle than P. sylvestris var. mongolica. In different water gradients, P. densiflora var. zhangwuensis had lower MDA contents, and higher CAT and Pro contents than P. sylvestris var. mongolica. In mild drought area(40%FMC, SWP=-4.82bar), MDA, CAT and Pro contents of P. sylvestris var. mongolica had a ascending trend in comparison with those in control area(100%FMC, SWP=-2.81bar), and the three physiological indicators had a further increase in moderate drought area(30%FMC, SWP=-10.35bar) and reached to the maximum in serious drought area(20%FMC, SWP=-20.14bar). By contrast in mild drought area these three physiological indicators in leaves of P. densiflora var. zhangwuensis were lower than those in control area, and began to increase in moderate drought area, especially the CAT content was significantly higher than that in control area, and they also reached to the maximum in serious drought area. The above results determinated that P. densiflora var. zhangwuensis showed stress symptoms at a lower soil moisture, and it could endure and adapt to drought stress by improving protective enzyme activity(CAT) and increasing osmotic substance(Pro). At different moisture gradients, both Chl and LWC of P. densiflora var. zhangwuensis were more than those of P. sylvestris var. mongolica, and these two indicators in the same species had the same changing trend,which was : CK > mild drought area > moderate drought area > serious drought area.
For an objective evaluation of salinity-alkalinity tolerance and a better understanding of its mechanisms in Pinus densiflora var. zhangwuensis, 4-year-old P. densiflora var. zhangwuensis and P. sylvestris var. mongolica were studied. Pot experiments were used to examine the effects of three types of salt (NaCl, Na2CO3, and NaHCO3) and alkali stress (NaOH) on seedling growth and physiological and biochemical indices. The substrate was fine sand taken from Zhanggutai sandy land, Liaoning Province, China. In the salt stress treatments, the matrix salt content was 4 g salt kg-1, and in NaOH stress treatments, the matrix pH value was 9.0. In P. densiflora var. zhangwuensis, the injury level was smaller, and the root tolerance index was bigger under salt-alkali stress. Malondialdehyde (MDA) content did not significantly increase under salt-alkali stress, while catalase (CAT) activity was significantly induced by stress, by as much as 22.6 times that of the control. Finally, chlorophyll (Chl) content did not significantly decline, and leaf water content (LWC) increased slightly. Pinus sylvestris var. mongolica responded differently to salt-alkali stress; its MDA content increased (NaCl treatment) or significantly increased (remaining three treatments), but its CAT activity did not increase significantly. Leaf Chl content declined significantly under salt-alkali stress, with Chl contents in the Na2CO3, NaHCO3, NaCl treatments 36.2%, 21.7%, and 62.3% lower than in the control. In addition, LWC decreased in leaves under stress. The results suggest that P. densiflora var. zhangwuensis has greater salt-alkali tolerance than P. sylvestris var. mongolica and that it ensures this tolerance by increasing CAT activity and maintaining high contents of Chl and leaf water. Higher levels of iron in needles of P. densiflora var. zhangwuensis enhanced CAT activity and Chl content, while higher levels of elemental zinc and copper were also associated with stronger resistance. In P. densiflora var. zhangwuensis, salinity stress did not cause drought stress, so growth was not inhibited by drought stress, but by ion toxicity and nutrient imbalance due to salinity stress.
Freezer method was used to carry out cold-stress test at different temperature gradients. With the combination between conductivity method and physiological method, relative conductivity as well as physiological and biochemical indicators such as MDA, CAT, Pro and SS in leaves of seedlings before and after stress were measured in order to identify cold resistance of the two species and to clarify their mechanism of cold hardiness. The results showed that: After cold resistance exercise, half-lethal temperature of P. densiflora var. zhangwuensis and P. sylvestris var. mongolica was -54.23℃and -50.34℃, respectively, which indicated that both species were strong cold hardiness, but a little cold hardiness in P. densiflora var. zhangwuensis higher than P. sylvestris var. mongolica due to cold acclimation. With the deepening of stress, Pro and SS changes model of two species was basically the same, while CAT activity in different patterns. In lower than -20℃, the CAT activity in leaves of P. densiflora var. zhangwuensis seedlings gradually increased, while that of P. sylvestris var. mongolica remained constant. In every temperature gradient, Pro and SS content was lower in leaves of P. densiflora var. zhangwuensis than P. sylvestris var. mongolica, while CAT activity of the former higher than the latter. The two species had slightly different cold-resistance mechanisms: From the point of view of content of indicators, P. densiflora var. zhangwuensis focused on using higher antioxidant enzyme activities to resist stress injury, and P. sylvestris var. mongolica focused on using higher ability of osmotic adjustment to enhance cold tolerance. From the point of view of variation trend in the indicators with the stress deepening, in mild cold stress, the two species could increase cold hardiness by increasing osmolyte (Pro and SS) content. But in severe cold stress (-40℃~-60℃) that caused stress injuries, P. densiflora var. zhangwuensis had a smaller increase of MDA by increasing CAT activity and maintaining the SS increments, while P. sylvestris var. mongolica had no such mechanism.
In order to photosynthesize, land plants must open their stomata to exchange small amounts of CO2 at the cost of losing a lot of water vapor through transpiration. This gas exchange is unequal in that more water is lost than CO2 is taken in. In general, fast growing species are characterized by their high photosynthetic capacity, most of which is achieved at the expense of consuming large amounts of water. This trade-off poses a serious challenge to forestry in the selection of fast-growing tree species in arid and semiarid areas. The ideal forestry species for drier climates should maintain a high photosynthetic capacity as well as low water consumption for transpiration, but such species are very rare. In this study, the LI-6400 (LICOR, Inc. Lincoln, NE, USA) portable photosynthesis system was employed to measure diurnal changes in photosynthetic and transpiration indices in sexually mature P. densiflora var. zhangwuensis and P. sylvestris var. mongolica (18 years old). In addition, the cut-branch transpiration method was used to compare stomatal and cuticular transpiration in order to evaluate the sensitivity of stomatal opening and closing and to characterize the photosynthetic productivity and water consumption for transpiration in these two species in different growing seasons. P. densiflora var. zhangwuensis had a higher photosynthesis rate (Pn) and a lower transpiration rate (Tr) than P. sylvestris var. mongolica under the same conditions. In May and July, the diurnal changes of Pn and Tr in P. densiflora var. zhangwuensis formed double-peaked curve as a result of mid-day stomatal limitation, while it formed single-peaked curve in October. The daily variation of Pn and Tr in P. sylvestris var. mongolica exhibited a monopeak curve throughout the growing season. The net photosynthesis per day in P. densiflora var. zhangwuensis was higher than in P. sylvestris var. mongolica; values for the former were 163.4 (May), 211.1 (July), and 183.6 (October) percent of the latter. The photoresponse parameter measurements showed that, in different months, the maximum rate of photosynthesis of P. densiflora var. zhangwuensis was greater than that of P. sylvestris var. mongolica, and the former also had a higher light saturation point and a lower light compensation point. The stomatal conductivity (Gs) and Tr of P. densiflora var. zhangwuensis were lower than those of P. sylvestris var. mongolica when measured at random times. The stomatal and cuticular Tr of P. densiflora var. zhangwuensis were lower, and in the same drought conditions, its stomata were deeper and had a higher sensitivity for opening and closing. Water use efficiency (WUE) of P. densiflora var. zhangwuensis was 2.29 times that of P. sylvestris var. mongolica. Analysis of the correlation between Gs and WUE showed that P. densiflora var. zhangwuensis could maintain high WUE when Gs levels were high. In May and July, when P. densiflora var. zhangwuensis exhibited midday photosynthetic depression, its Gs level was low, usually in the 90~200 mmol ? m-2 ? s-1 range. At the same time, its WUE was higher than that of P. sylvestris var. mongolica, indicating that P. densiflora var. zhangwuensis could effectively conserve water by closing its stomata rapidly at midday, so as to maintain its high WUE. Nitrogen content per unit leaf area(Narea) and photosynthetic nitrogen use efficiency (PNUE) was higher, and gaseous conductance through intercellular air space(gias) was also larger in P. densiflora var. zhangwuensis. The stomata diameter and density was small, which was also able to maintain a smaller Gs and Tr, and thus a reason to maintain a larger WUE in P. densiflora var. zhangwuensis. These results implied that P. densiflora var. zhangwuensis can improve its WUE, yielding its higher Pn and lower Tr, and achieves its fast growth in arid and semiarid regions through its sensitive stomatal response and leaf xeromorphism.
引文
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