遮光与淹水胁迫对白栎生理特性的影响
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摘要
本文以白栎(Quercus fabri)为研究对象,分别对2年生白栎实生苗进行遮光处理(设4个光照水平：全光照、30%遮光、60%遮光、80%遮光)和对1年生白栎实生苗进行淹水胁迫处理(设4个土壤水分水平,CK水平：对照水平,土壤含水量为田间持水量的75%左右；S1水平：水面在土壤表面以下15cm；S2水平：土壤含水量处于完全饱和状态；S3水平：土壤含水量过饱和,水面高于土面4cm左右),处理一定时间后对植株叶片的生理指标、光合特性进行研究,探讨白栎在不同光照强度以及淹水胁迫下的生理反应及适应性,为白栎栽培提供理论依据。本研究结果如下：
(1)随着遮光程度的增加,白栎叶片叶绿素a、叶绿素b、总叶绿素含量升高。相关分析表明,叶绿素总量与可溶性糖含量成极显著负相关,与叶绿素a、叶绿素b和蛋白质成极显著正相关,与淀粉和叶片相对含水量无相关性。
(2)4种光照强度下白栎净光合速率(Pn)日变化曲线为单峰型,遮光处理降低了白栎叶片净光合速率(Pn)、蒸腾速率(Tr)和气孔导度(Gs)；叶片胞间CO2浓度(Ci)在200-400μmol·mol-1之间波动；随着遮光程度的增加光补偿点(LCP)、CO2补偿点(CCP)升高,光饱和点(LSP)、CO2饱和点(CSP)、表观量子效率(AQY)、羧化效率(CE)、光饱和时最大净光合速率(Pn)和CO2饱和时最大净光合速率(Pn)均降低,全光照处理下CO2饱和点最高,为1320μmol·mol-1,80%遮光处理下CO2补偿点最高,为48.69μmol·mol-1,30%遮光处理下CO2补偿点最低,为23.43μmol·mol-1.
(3)对不同光照条件下白栎叶片叶绿素a荧光参数进行测定。结果表明,随着遮光程度的增加,PSⅡ原初光能转化效率(Fv/Fm)、潜在活性(Fv/Fo)值增加,光化学淬灭(qP)升高,非光化学淬灭(qN)降低；表观光合电子传递速率(ETR)全光照>30%>60%遮光>80%遮光。不同遮光处理有限地提高了白栎叶片PSⅡ吸收光能的利用效率,60%遮光度以下白栎能通过自身调节适应弱光环境,具有一定的耐荫性,但严重遮光(80%遮光)时光合速率下降较多,不利于其正常生长。
(4)随着淹水胁迫程度的增加,白栎叶片叶绿素a、叶绿素b、总叶绿素含量降低。S3水平处理21天后白栎叶片失绿,胁迫35天后失绿现象严重,50%植株死亡。对各生理指标相关分析表明,叶绿素总量与可溶性糖含量、丙二醛含量、叶片相对含水量成极显著负相关,与电导率成显著负相关,与叶绿素a、叶绿素b成极显著正相关,与蛋白质含量无相关性。
(5)淹水胁迫降低了白栎净光合速率和气孔导度,提高了水分利用效率。4种水平下白栎净光合速率日变化趋势基本一致,呈“单峰”曲线；气孔导度在胁迫前后变化趋势相同,为上升-下降趋势,呈单峰型；水分利用效率日变化呈近“L”型,最高值均出现在早8：00。
(6)随着淹水胁迫程度的增加,白栎光补偿点、CO2补偿点升高；光饱和点、C02饱和点、表观量子效率、羧化效率、光饱和时最大净光合速率和C02饱和时最大净光合速率均降低；S3水平处理下PSⅡ原初光能转化效率(Fv/Fm)和潜在活性(Fv/Fo)值最低,说明S3水平处理下光合电子传递效率最低,使白栎叶片反映中心捕获激发能效率降低,不利于植物进行光合作用。白栎在重度淹水胁迫下会使叶片失绿,降低叶片光能转换效率,失去一定的观赏效果,但植株本身具有很强的自我调节能力,能迅速调节自身的新陈代谢,作出生理响应,具有较强的耐涝性。
In the present work, the Quercus fabri were studied. The biennial seedlings of Quercus fabri were under 0%,30%,60% and 80% shading treatments and the annual seedlings of Quercus fabri were dealed with by submergence stress. Four levels of soil water content named CK, S1,S2 and S3 were set up in this study, where CK means the soil moisture was about 75% of field capacity, S1 means water surface under 15 cm of the soil surface, S2 means the soil moisture was under saturation state and S3 means the soil moisture was under super-saturation state and water surface was 15 cm higher than the soil surface. The physiological indexes and photosynthetic characteristics of leaves were studied after the shading treatment (biennial seedlings of Quercus fabri) and submergence stress (annual seedlings of Quercus fabri). The physiological response and adaptability of Quercus fabri under different light and water conditions were discussed. The results would provide a theory basis for the cultivation of Quercus fabri. The main conclusions as followed:
(1)The content of chlorophyll a, chlorophyll b, total chlorophyll increased with the degree of shading treament. A negative correlation was found between the content of total chlorophyll and soluble sugar. The content of total chlorophyll was positive correlated with chlorophyll a, chlorophyll b and protens. Besides that, there was no obvious correlation between the starch, leaf relative water and total chlorophyll content.
(2) The diurnal variations of net photosynthetic rate (Pn) for Quercus fabri showed single peak curves under four light intensity. The shading treatment resulted in the decrease of net photosynthetic rate (Pn), transpiration rate (Tr) and stomatal conductance(Gs) of Quercus fabri. The intercellular CO2 concentration (Ci) of leaves was ranging from 200 to 400μmol·mol-1. Except the light compensation points (LCP) and CO2 compensation(CCP) points, the light saturation points (LSP),CO2 saturation points (CSP), apparent quantum yield (AQY), carboxylation efficiency (CE) and the maximal net photosynthetic rates (Pn) under light and CO2 saturation conditions all decreased with the shading treatment. The maximum value of CSP and CCP was 1320μmol·mol-1 and 48.69μol·mol-1 under 0% and 80% shading treatment, respectively. The minimum value of CCP was 23.43μmol·mol-1 under 30% shading treatment.
(3) The chlorophyll a fluorescence parameters of Quercus fabri were determined under different light conditions.The results showed that, the PSII primary chemical efficiency (Fv/Fm), potential activity (Fv/Fo) and photochemical quenching (qP) increased and non-photochemical quenching (qN) decreased with the shading treatment. The apparent photosynthetic electron transport rate (ETR):0%>30%>60%>80% shading treatment. The different shading treatments improved the utilization efficiency of PS II absorbing light energy in some extent. Quercus fabri has shade tolerance ability and can adjust itself to a weak light environment. The photosynthesis rate decreased greatly under 80% shading treatment which is very unfavorable to growth.
(4) The content of chlorophyll a, chlorophyll b, total chlorophyll decreased with the degree of submergence stress.Under the S3 stress, the leaves of Quercus fabri changed into yellow gradually after 21 days and the situation became more serious after 35 days,50% plants have died. The results for the studies on physiological indexes showed that, there were significant negative correlation between the total chlorophyll content and soluble sugar, malondialdehyde and the leaf relative water. In addition, the negative correlation between total chlorophyll content and conductivity also obvious. On the contrary, the total chlorophyll content were remarkable positive correlated to chlorophyll a and chlorophyll b. There was no correlation between proten and the total chlorophyll content.
(5) The net photosynthetic rate and stomatal conductance of Quercus fabri were decreased and water utilization efficiency was increased with the submergence stress, respectively. The diurnal trends of net photosynthetic rate under these 4 stress situations were basically consistent and all showed single peak curve. The variation of stomatal conductance before and after the submergence stress also very similar, which showed ascending-descending trend. The water utilization efficiency showed "L" trend with the peak value appeared at 8:00 AM.
(6) The light compensation points and CO2 compensation points were increased and the light saturation points, CO2 saturation points, apparent quantum yield, carboxylation efficiency and the maximal net photosynthetic rate under light and CO2 saturation conditions were decreased with the submergence stress. The PSII primary chemical efficiency (Fv/Fm) and potential activity (Fv/Fo) were lowest under S3 level, which demonstrated that the photosynthetic electron transport rate was lowest. Meanwhile, the capture efficiency of reaction center to excited energy was descreased, which is an unfavorable condition to the photosynthesizes of Quercus fabri. The severe submergence stress caused the leaves chlorosis, decreased the light energy transform efficiency and made the loss of certain ornamental effect. However, the plant of Quercus fabri has a strong waterlogging tolerance for its self-adjusting function and can quickly adjust its metabolism to physiological response.
(1)随着遮光程度的增加,白栎叶片叶绿素a、叶绿素b、总叶绿素含量升高。相关分析表明,叶绿素总量与可溶性糖含量成极显著负相关,与叶绿素a、叶绿素b和蛋白质成极显著正相关,与淀粉和叶片相对含水量无相关性。
(2)4种光照强度下白栎净光合速率(Pn)日变化曲线为单峰型,遮光处理降低了白栎叶片净光合速率(Pn)、蒸腾速率(Tr)和气孔导度(Gs)；叶片胞间CO2浓度(Ci)在200-400μmol·mol-1之间波动；随着遮光程度的增加光补偿点(LCP)、CO2补偿点(CCP)升高,光饱和点(LSP)、CO2饱和点(CSP)、表观量子效率(AQY)、羧化效率(CE)、光饱和时最大净光合速率(Pn)和CO2饱和时最大净光合速率(Pn)均降低,全光照处理下CO2饱和点最高,为1320μmol·mol-1,80%遮光处理下CO2补偿点最高,为48.69μmol·mol-1,30%遮光处理下CO2补偿点最低,为23.43μmol·mol-1.
(3)对不同光照条件下白栎叶片叶绿素a荧光参数进行测定。结果表明,随着遮光程度的增加,PSⅡ原初光能转化效率(Fv/Fm)、潜在活性(Fv/Fo)值增加,光化学淬灭(qP)升高,非光化学淬灭(qN)降低；表观光合电子传递速率(ETR)全光照>30%>60%遮光>80%遮光。不同遮光处理有限地提高了白栎叶片PSⅡ吸收光能的利用效率,60%遮光度以下白栎能通过自身调节适应弱光环境,具有一定的耐荫性,但严重遮光(80%遮光)时光合速率下降较多,不利于其正常生长。
(4)随着淹水胁迫程度的增加,白栎叶片叶绿素a、叶绿素b、总叶绿素含量降低。S3水平处理21天后白栎叶片失绿,胁迫35天后失绿现象严重,50%植株死亡。对各生理指标相关分析表明,叶绿素总量与可溶性糖含量、丙二醛含量、叶片相对含水量成极显著负相关,与电导率成显著负相关,与叶绿素a、叶绿素b成极显著正相关,与蛋白质含量无相关性。
(5)淹水胁迫降低了白栎净光合速率和气孔导度,提高了水分利用效率。4种水平下白栎净光合速率日变化趋势基本一致,呈“单峰”曲线；气孔导度在胁迫前后变化趋势相同,为上升-下降趋势,呈单峰型；水分利用效率日变化呈近“L”型,最高值均出现在早8：00。
(6)随着淹水胁迫程度的增加,白栎光补偿点、CO2补偿点升高；光饱和点、C02饱和点、表观量子效率、羧化效率、光饱和时最大净光合速率和C02饱和时最大净光合速率均降低；S3水平处理下PSⅡ原初光能转化效率(Fv/Fm)和潜在活性(Fv/Fo)值最低,说明S3水平处理下光合电子传递效率最低,使白栎叶片反映中心捕获激发能效率降低,不利于植物进行光合作用。白栎在重度淹水胁迫下会使叶片失绿,降低叶片光能转换效率,失去一定的观赏效果,但植株本身具有很强的自我调节能力,能迅速调节自身的新陈代谢,作出生理响应,具有较强的耐涝性。
In the present work, the Quercus fabri were studied. The biennial seedlings of Quercus fabri were under 0%,30%,60% and 80% shading treatments and the annual seedlings of Quercus fabri were dealed with by submergence stress. Four levels of soil water content named CK, S1,S2 and S3 were set up in this study, where CK means the soil moisture was about 75% of field capacity, S1 means water surface under 15 cm of the soil surface, S2 means the soil moisture was under saturation state and S3 means the soil moisture was under super-saturation state and water surface was 15 cm higher than the soil surface. The physiological indexes and photosynthetic characteristics of leaves were studied after the shading treatment (biennial seedlings of Quercus fabri) and submergence stress (annual seedlings of Quercus fabri). The physiological response and adaptability of Quercus fabri under different light and water conditions were discussed. The results would provide a theory basis for the cultivation of Quercus fabri. The main conclusions as followed:
(1)The content of chlorophyll a, chlorophyll b, total chlorophyll increased with the degree of shading treament. A negative correlation was found between the content of total chlorophyll and soluble sugar. The content of total chlorophyll was positive correlated with chlorophyll a, chlorophyll b and protens. Besides that, there was no obvious correlation between the starch, leaf relative water and total chlorophyll content.
(2) The diurnal variations of net photosynthetic rate (Pn) for Quercus fabri showed single peak curves under four light intensity. The shading treatment resulted in the decrease of net photosynthetic rate (Pn), transpiration rate (Tr) and stomatal conductance(Gs) of Quercus fabri. The intercellular CO2 concentration (Ci) of leaves was ranging from 200 to 400μmol·mol-1. Except the light compensation points (LCP) and CO2 compensation(CCP) points, the light saturation points (LSP),CO2 saturation points (CSP), apparent quantum yield (AQY), carboxylation efficiency (CE) and the maximal net photosynthetic rates (Pn) under light and CO2 saturation conditions all decreased with the shading treatment. The maximum value of CSP and CCP was 1320μmol·mol-1 and 48.69μol·mol-1 under 0% and 80% shading treatment, respectively. The minimum value of CCP was 23.43μmol·mol-1 under 30% shading treatment.
(3) The chlorophyll a fluorescence parameters of Quercus fabri were determined under different light conditions.The results showed that, the PSII primary chemical efficiency (Fv/Fm), potential activity (Fv/Fo) and photochemical quenching (qP) increased and non-photochemical quenching (qN) decreased with the shading treatment. The apparent photosynthetic electron transport rate (ETR):0%>30%>60%>80% shading treatment. The different shading treatments improved the utilization efficiency of PS II absorbing light energy in some extent. Quercus fabri has shade tolerance ability and can adjust itself to a weak light environment. The photosynthesis rate decreased greatly under 80% shading treatment which is very unfavorable to growth.
(4) The content of chlorophyll a, chlorophyll b, total chlorophyll decreased with the degree of submergence stress.Under the S3 stress, the leaves of Quercus fabri changed into yellow gradually after 21 days and the situation became more serious after 35 days,50% plants have died. The results for the studies on physiological indexes showed that, there were significant negative correlation between the total chlorophyll content and soluble sugar, malondialdehyde and the leaf relative water. In addition, the negative correlation between total chlorophyll content and conductivity also obvious. On the contrary, the total chlorophyll content were remarkable positive correlated to chlorophyll a and chlorophyll b. There was no correlation between proten and the total chlorophyll content.
(5) The net photosynthetic rate and stomatal conductance of Quercus fabri were decreased and water utilization efficiency was increased with the submergence stress, respectively. The diurnal trends of net photosynthetic rate under these 4 stress situations were basically consistent and all showed single peak curve. The variation of stomatal conductance before and after the submergence stress also very similar, which showed ascending-descending trend. The water utilization efficiency showed "L" trend with the peak value appeared at 8:00 AM.
(6) The light compensation points and CO2 compensation points were increased and the light saturation points, CO2 saturation points, apparent quantum yield, carboxylation efficiency and the maximal net photosynthetic rate under light and CO2 saturation conditions were decreased with the submergence stress. The PSII primary chemical efficiency (Fv/Fm) and potential activity (Fv/Fo) were lowest under S3 level, which demonstrated that the photosynthetic electron transport rate was lowest. Meanwhile, the capture efficiency of reaction center to excited energy was descreased, which is an unfavorable condition to the photosynthesizes of Quercus fabri. The severe submergence stress caused the leaves chlorosis, decreased the light energy transform efficiency and made the loss of certain ornamental effect. However, the plant of Quercus fabri has a strong waterlogging tolerance for its self-adjusting function and can quickly adjust its metabolism to physiological response.
引文
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