Chlorophyll fluorescence characteristics of <i>Phyllostachys edulis</i> during its fast growth period

ZHOU Zheyu; XU Chao; HU Ce; WANG Haixiang; LIANG Xieen; ZHANG Rumin; WEN Guosheng

doi:10.11833/j.issn.2095-0756.2018.01.010

Volume 35 Issue 1

Jan. 2018

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ZHOU Zheyu, XU Chao, HU Ce, WANG Haixiang, LIANG Xieen, ZHANG Rumin, WEN Guosheng. Chlorophyll fluorescence characteristics of Phyllostachys edulis during its fast growth period[J]. Journal of Zhejiang A&F University, 2018, 35(1): 75-80. doi: 10.11833/j.issn.2095-0756.2018.01.010

Citation:

ZHOU Zheyu, XU Chao, HU Ce, WANG Haixiang, LIANG Xieen, ZHANG Rumin, WEN Guosheng. Chlorophyll fluorescence characteristics of Phyllostachys edulis during its fast growth period[J]. Journal of Zhejiang A&F University, 2018, 35(1): 75-80. doi: 10.11833/j.issn.2095-0756.2018.01.010

Chlorophyll fluorescence characteristics of Phyllostachys edulis during its fast growth period

doi: 10.11833/j.issn.2095-0756.2018.01.010

1.
School of Forestry and Biotechnology, Zhejiang A & F University, Hangzhou 311300, Zhejiang, China
2.
State Key Laboratory of Subtropical Silviculture, Zhejiang A & F University, Hangzhou 311300, Zhejiang, China

Received Date: 2016-11-15
Rev Recd Date: 2017-01-08
Publish Date: 2018-02-20

Abstract

To explore chlorophyll fluorescence characteristics of a Phyllostachys edulis blade during its fast growth period (early, middle, and late phases), parameters concerning chlorophyll fluorescence of a Ph. edulis blade were measured using PAM-2100 chlorophyll fluorescence spectrometer during its fast growth period of the Bamboo Physiological and Ecological Monitoring Station of Zhejiang A & F University, also comprehensive evaluation of its fluorescence characteristics was then conducted. The results showed that the actual photochemical quantum yield of Photosystem Ⅱ (PS-Ⅱ), the maximal photochemical efficiency of PS-Ⅱ (F_v/F_m), and the potential PS-Ⅱ activity for same-aged Ph. edulis samples had significant differences within different stages of the rapid growth period (P < 0.05) going from a lower level to a higher level for the early phase, the middle phase, and the latter phase. Variation in the F_v/F_m ratio was 0.672-0.773 and in the F_v /F_o ratio was 2.068-3.231. During the fast grwoth period, the value of Non-photochemical Quenching(q_NP) in the order: middle phase < late phase < early phase. The q_NP value also showed strong daily variation during different phases of the fast growth period. Chlorophyll fluorescence parameters of bamboo blade show significant changes during its fast growth period. Thus, as indicated by the comprehensive comparison, photosynthesis of the Ph. edulis blade showed an increasing tendency during its rapid growth.
- botany,
- Phyllostachys edulis,
- fast growth period,
- chlorophyll fluorescence

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叶绿素荧光与光合作用中各个反应过程密切相关，植物体内的叶绿素荧光信号能够快速灵敏地反映植物光合生理状况。与“表观性”的气体交换指标相比，叶绿素荧光参数更具“内在性”的特点^[1-3]。目前，植物叶片叶绿素荧光特性已经被广泛应用到植物光合机制和逆境生理等研究领域^[4-5]。毛竹Phyllostachys edulis属禾本科Gramineae植物，广泛分布于中国南方亚热带地区。中国毛竹林面积已达270万hm²，是中国竹类植物中分布最广、面积最大的竹种^[6]。毛竹有着与其他树种不同的生长方式即“爆发式生长”^[7]。“爆发式生长”又称快速生长，指毛竹出笋后的伸长生长，从3月底开始约60 d内，从笋长成10 m以上的竹秆。为探讨毛竹快速生长的机制，从毛竹不同生长期的蒸腾速率、气孔导度、水分利用关系、器官变化、水势变化及液流特征和环境因子的关系等方面进行了研究^[8-11]，但是针对毛竹快速生长期的叶片叶绿素荧光参数的研究较为匮乏。本研究测定了毛竹快速生长不同时期叶片叶绿素荧光参数，分析其变化规律，试图探究毛竹快速生长与毛竹叶片叶绿素荧光参数之间的关系，为揭示毛竹快速生长机制提供一定的理论依据。

1. 材料与方法

1.1. 实验地概况与实验材料

实验地位于浙江省杭州市临安区东部的青山镇研里村(30°14′N，119°42′E)的浙江农林大学毛竹生理生态监测定位站。实验地地处低山丘陵区，海拔为48~50 m。属中亚热带季风气候，光照充足，雨水充沛。年平均气温为15.9 ℃，全年降水量为1 427.0 mm，全年日照时数为1 920.0 h。选取实验地竹林向阳方向，长势良好的1年生和2年生毛竹各2株，搭建观测塔，选取林冠中层健康完好的叶片测定叶绿素荧光参数。

1.2. 实验方法

1.2.1. 毛竹年龄的确定

通过对毛竹外部特征(枝痕个数、基部笋箨、皮色、白粉环和附着物等)的观察，并结合当地竹农的意见，确定毛竹年龄进行，作标记。

1.2.2. 叶绿素荧光的测定

从2016年3月底开始，每月选择天气晴朗，光照较好的一天，利用PAM-2100型荧光仪对选取的实验地毛竹叶片进行叶绿素荧光测定。在8：30-16：30，2 h测定1次，重复4次·株^-1。叶片经过30 min暗适应后测定初始荧光产量(F_o)，最大荧光产量(F_m)，最大光化学量子产量光系统Ⅱ(PSⅡ)的最大光化学效率(F_v/F_m)。在自然光下适应30 min，当荧光基本稳定时测定实际光化学量子产量(Y_ield)，光下最大荧光(F_m′)。计算出可变荧光(F_v)，PSⅡ潜在光化学效率(F_v/F_o)，非光化学猝灭系数(q_NP)。

1.3. 数据处理

取2016年3月25日，4月18日，5月11日的毛竹叶片叶绿素荧光数据分别作为毛竹快速生长前期、中期、后期的叶片叶绿素荧光数据。利用SPSS 19.0对不同生长期不同竹龄的毛竹叶绿素荧光参数进行单因素方差分析(one-way ANOVA)，使用最小显著性差异(LSD)法进行多重比较。用Excel制图。

2. 结果与分析

2.1. 不同生长期不同竹龄毛竹叶片F_v/F_m日变化

F_v/F_m指最大光化学效率，是用来研究植物逆境响应的重要参数，主要用于判断植物是否受到了光抑制。F_v/F_m在非胁迫下变化极小，且不受物种和生长条件影响，在受到胁迫的情况下F_v/F_m显著降低^[7]。正常情况下植物叶片F_v/F_m的数值为0.75~0.85^[12]。

不同生长期不同竹龄毛竹的F_v/F_m如表 1所示。在同一生长期内，毛竹叶片的F_v/F_m测定值无明显日变化。毛竹叶片在快速生长的前期和中期F_v/F_m测定值均低于0.75，表明在这一阶段毛竹叶片受到了环境胁迫。在同一生长期内，不同竹龄毛竹叶片的F_v/F_m测定值差异显著。不同生长期毛竹叶片的F_v/F_m测定值均为1年生毛竹＜2年生毛竹。表明2年生毛竹叶片在快速生长期内有更高的光能利用率。不同生长期内，同一竹龄毛竹叶片的F_v/F_m测定值均为前期＜中期＜后期。不同生长期对同一竹龄毛竹叶片F_v/F_m影响达到了显著水平(P＜0.05)。反映在整个生长期内，毛竹叶片光能利用率后期最强，中期居中，前期最弱，2年生毛竹大于1年生毛竹。

测定时间	前期		中期		后期
测定时间	1年生	2年生	1年生	2年生	1年生	2年生
8:30	0.675 ± 0.011 bc	0.691 ± 0.008 ac	0.693 ± 0.017 bb	0.722 ± 0.044 ab	0.740 ± 0.015 ba	0.759 ± 0.014 aa
10:30	0.678 ± 0.027 bc	0.684 ± 0.013 ac	0.711 ± 0.022 bb	0.739 ± 0.028 ab	0.743 ± 0.017 ba	0.761 ± 0.005 aa
12:30	0.679 ± 0.006 bc	0.695 ± 0.039 ac	0.703 ± 0.030 bb	0.726 ± 0.010 ab	0.756 ± 0.003 ba	0.773 ± 0.009 aa
14:30	0.672 ± 0.010 bc	0.684 ± 0.005 ac	0.708 ± 0.017 bb	0.732 ± 0.013 ab	0.760 ± 0.013 ba	0.771 ± 0.021 aa
16:30	0.676 ± 0.016 bc	0.693 ± 0.008 ac	0.707 ± 0.026 bb	0.729 ± 0.021 ab	0.750 ± 0.008 ba	0.768 ± 0.016 aa
说明:数字后第1个小写字母不相同代表同一生长期内不同竹龄之间差异显著, 第2个小写字母不相同代表不同生长期内同一竹龄之间差异显著

Table 1. Diurnal changes of F_v/F_m of Phyllostachys edulis in different growth stages

2.2. 不同生长期不同竹龄毛竹叶片F_v/F_o日变化

F_o表示PSⅡ反应中心全部开放式即原初电子受体全部氧化时的荧光水平，理论上用来指反应中心未能发生光化学反应时的叶绿素荧光。F_v/F_o常用来度量PSⅡ的潜在光化学效率，它对环境引起的变化比F_v/F_m更加敏感。毛竹快速生长期不同时期的F_v/F_o测定值见表 2。在毛竹快速生长期的不同时期，毛竹叶片F_v/F_o值无明显日变化，表明在同一时期实验区的环境较为稳定无明显变化。

测定时间	前期		中期		后期
测定时间	1年生	2年生	1年生	2年生	1年生	2年生
8:30	2.068 ± 0.128 bc	2.312 ± 0.105 ac	2.254 ± 0.148 bb	2.489 ± 0.351 ab	2.671 ± 0.142 ba	2.984 ± 0.140 aa
10:30	2.121 ± 0.166 bc	2.178 ± 0.136 ac	2.377 ± 0.186 bb	2.701 ± 0.314 ab	2.788 ± 0.189 ba	3.019 ± 0.073 aa
12:30	2.144 ± 0.085 bc	2.349 ± 0.249 ac	2.301 ± 0.225 bb	2.533 ± 0.123 ab	3.121 ± 0.056 ba	3.231 ± 0.105 aa
14:30	2.099 ± 0.119 bc	2.181 ± 0.071 ac	2.378 ± 0.142 bb	2.657 ± 0.141 ab	3.044 ± 0.135 ba	3.205 ± 0.246 aa
16:30	2.146 ± 0.144 bc	2.357 ± 0.114 ac	2.356 ± 0.229 bb	2.588 ± 0.246 ab	2.916 ± 0.107 ba	3.144 ± 0.151 aa
说明：数字后第1个小写字母不相同代表同一生长期内不同竹龄之间差异显著，第2个小写字母不相同代表不同生长期内同一竹龄之间差异显著。

Table 2. Diurnal changes of F_v/F_o of Phyllostcwhys edulis in different growth stages

在不同生长期内，毛竹叶片的F_v/F_o的差异均达到显著水平(P＜0.05)，呈现为前期＜中期＜后期。在同一生长期内，不同竹龄毛竹叶片F_v/F_o差异显著。在毛竹快速生长的前期、中期、后期，2年生毛竹叶片F_v/F_o均大于1年生毛竹。表明2年生毛竹叶片对环境胁迫有着更强的适应性。

2.3. 不同生长期不同竹龄毛竹叶片Y_ield日变化

实际光化学量子产量(Y_ield)是指植物光合作用下PSⅡ总的光化学量子产量，它一定程度上反映了PSⅡ反应中心在部分关闭情况下的实际原初光能捕获效率^[10-11]。较高的Y_ield值代表了更高的光能转换效率，能够为植物暗反应的光合碳同化积累更多的能量，促进碳同化的高效运转和有机物的积累^[13]。

从表 3可以看出：不同生长期毛竹叶片的Y_ield值，从8：30开始随着时间推移不断降低，在14：30达到最低点并开始回升。多重比较表明，2种竹龄的毛竹叶片在毛竹快速生长的不同时期，其实际光化学量子产量存在显著差异(P＜0.05)。在同一生长期中，2年生毛竹的Y_ield值均大于1年生毛竹，表明2年生毛竹的光合转化率要优于1年生的毛竹。在毛竹快速生长的不同阶段，不同竹龄毛竹的Y_ield值均为前期＜中期＜后期，反映了毛竹叶片光能利用率随着毛竹的快速生长而提高。这变化规律与毛竹F_v/F_m和F_v/F_o基本保持一致。

测定时间	前期		中期		后期
测定时间	1年生	2年生	1年生	2年生	1年生	2年生
8:30	0.416 ± 0.021 bc	0.433 ± 0.008 ac	0.565 ± 0.023 bb	0.585 ± 0.017 ab	0.598 ± 0.019 ba	0.617 ± 0.023 aa
10:30	0.402 ± 0.011 bc	0.426 ± 0.016 ac	0.524 ± 0.017 bb	0.557 ± 0.013 ab	0.567 ± 0.029 ba	0.591 ± 0.013 aa
12:30	0.371 ± 0.013 bc	0.393 ± 0.012 ac	0.484 ± 0.011 bb	0.500 ± 0.012 ab	0.535 ± 0.017 ba	0.551 ± 0.016 aa
14:30	0.357 ± 0.009 bc	0.375 ± 0.015 ac	0.469 ± 0.014 bb	0.479 ± 0.015 ab	0.513 ± 0.008 ba	0.538 ± 0.015 aa
16:30	0.389 ± 0.015 bc	0.428 ± 0.005 ac	0.485 ± 0.012 bb	0.508 ± 0.020 ab	0.525 ± 0.013 ba	0.558 ± 0.007 aa
说明：数字后第1个小写字母不相同代表同一生长期内不同竹龄之间差异显著，第2个小写字母不相同代表不同生长期内同一竹龄之间差异显著。

Table 3. Diurnal changes of y_ield of Phyllostachys edulis in different growth stages

2.4. 不同生长期不同竹龄毛竹叶片q_NP日变化

非光化学猝灭系数(q_NP)反映PSⅡ吸收的光能中不能用于光合电子传递而是以热的形式耗散掉的光能部分，是表示热耗散多少的指标^[14]。

不同生长期不同竹龄毛竹叶片q_NP日变化见表 4。毛竹不同生长期内，叶片q_NP从8：30开始随着时间推移不断升高，峰值出现在12：30至14：30，之后逐渐下降。这主要是因为随着光照强度的增加，叶片把吸收的光能较多地转化到热耗散，用于光化学反应上的能量随之减少。

测定时间	前期		中期		后期
测定时间	1年生	2年生	1年生	2年生	1年生	2年生
8:30	1.769 ± 0.102 c	1.745 ± 0.088 c	1.492 ± 0.069 a	1.514 ± 0.055 a	1.693 ± 0.101 b	1.705 ± 0.073 b
10:30	1.835 ± 0.099 c	1.833 ± 0.096 c	1.594 ± 0.058 a	1.602 ± 0.056 a	1.833 ± 0.059 b	1.845 ± 0.076 b
12:30	2.011 ± 0.083 c	1.997 ± 0.112 c	1.728 ± 0.081 a	1.745 ± 0.062 a	1.887 ± 0.067 b	1.916 ± 0.081 b
14:30	2.015 ± 0.097 c	1.991 ± 0.056 c	1.766 ± 0.078 a	1.779 ± 0.073 a	1.893 ± 0.092 b	1.922 ± 0.084 b
16:30	1.879 ± 0.075 c	1.871 ± 0.081 c	1.630 ± 0.042 a	1.641 ± 0.050 a	1.750 ± 0.063 b	1.801 ± 0.091 b
说明：数字后小写字母不相同代表不同生长期内同一竹龄之间差异显著。

Table 4. Diurnal changes of q_NP of Phyllostachys edulis in different growth stages

在同一生长期内，不同竹龄的毛竹q_NP差异不显著。在毛竹快速生长的前期、中期、后期，毛竹叶片q_NP呈现出先下降后上升的规律，不同竹龄毛竹叶片q_NP均为中期＜后期＜前期。这说明在毛竹快速生长的中期，毛竹叶片能更充分地利用吸收的光能进行光合作用，光合利用率更高。

3. 结论与讨论

通过叶绿素荧光参数可以了解毛竹光合作用的内在特征，F_v/F_m，F_v/F_o，Y_ield这3个参数一般被认为是判断植物光合作用大小的重要依据。张其德等^[15]研究表明：这3个参数之间具有很好的一致性。本研究表明：在毛竹快速生长的不同时期，毛竹叶片叶绿素荧光参数变化明显，随着毛竹的快速生长，不同竹龄的毛竹叶片F_v/F_m，F_v/F_o，Y_ield均显著上升，显示毛竹PSⅡ反应中心的开放程度、PSⅡ潜在光化学效率和最大光化学效率均得到了提升。由此可以判断：毛竹快速生长的不同时期，毛竹叶片的光合作用能力为前期＜中期＜后期。毛竹快速生长前期，由于气温较低，毛竹叶片易受到环境胁迫，低温对毛竹叶片PSⅡ反应中心内禀光能转化效率产生了可逆性的胁迫影响，使毛竹叶片F_v/F_m低于正常值，光化学效率降低^[16]。毛竹快速生长的中期，后期，气温回升，低温胁迫消失，毛竹叶片PSⅡ反应中心内禀光能转化效率得到了显著的提升，毛竹的光合作用能力增强。实验结果显示：PSⅡ潜在活性(F_v/F_o)和F_v/F_m的变化规律基本一致，推测环境因素在这一阶段对毛竹的光合作用能力有着显著影响。

对不同竹龄毛竹的叶绿素荧光参数研究发现，2年生竹比1年生竹有着更强的光合作用能力。这是因为2年生竹需要为幼竹的生长提供大量的有机物质。叶淑贤等^[17]和施建敏等^[18]对毛竹光合作用季节变化规律进行了研究，证明在快速生长过程中老竹会为新竹提供一部分生长所需的营养物质。说明2年生竹为了满足新竹的需要，加强了其光合作用效率。

在毛竹快速生长期的不同时期，非光化学猝灭系数(q_NP)先降低后升高。植物叶片在吸收光能之后一般有3种利用途径：一种是以热量耗散的部分，一种是用于光化学反应的部分，还有一种是反应中心由非光化学反应耗散的能量^[19]，光合作用和热耗散的变化会引起相应的荧光变化^[20]。在毛竹快速生长初期，毛竹光合作用能力较弱，吸收的光能更多被用于热耗散，因此q_NP较高。随着毛竹的快速生长，毛竹光合作用能力增强，光能转化效率提升，q_NP也因此降低。在毛竹快速生长的后期，因为气温升高，光照强度增强，植物通过增加对吸收光能的利用和耗散避免过量光对光合机构造成伤害^[21]，因此毛竹的q_NP上升。

结合张守仁^[2]和温国胜等^[3]对叶绿素荧光参数的研究，证明叶绿素荧光参数反映了植物的光合作用能力。毛竹的快速生长和其本身的光合作用能力有着密切的关系，这种关系在毛竹快速生长的中后期尤为明显。毛竹快速生长的中后期是毛竹生长最快的时期，可以认为毛竹光合作用能力的增强是毛竹快速生长的重要因素。

本研究仅对1~2年生毛竹的叶绿素荧光参数进行了研究。今后可以进一步从环境因素、毛竹竹龄等影响因素来探求它们和毛竹叶绿素荧光参数之间的深层联系，进一步深入分析毛竹快速生长的生理机制。

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Chlorophyll fluorescence characteristics of Phyllostachys edulis during its fast growth period

doi: 10.11833/j.issn.2095-0756.2018.01.010