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三叶青Tetrastigma hemsleyanum是葡萄科Vitaceae崖爬藤属Tetrastigma的一种珍稀中草药[1],具有抗炎、镇痛等作用[2],临床上常用来抑制结肠癌[3]、食管癌[4]、肺癌[5-6]等多种肿瘤细胞的增殖和迁移。因其药理作用明显,致使采挖量逐年增加,林下野生三叶青濒临灭绝。近年来,为满足医疗需求,人工栽培面积不断扩大。三叶青在全光照下生长不良或死亡,探寻其适宜生长的光照条件势在必行。在药用植物生长过程中,光强影响其光合作用,进而影响其质量和产量[7]。万小燕[8]对3种药用植物进行了遮光处理,发现遮光影响凉粉草Mesona chinensis形态特征与光合性能,适当遮光可以提高三甲仙草Mesona paruifsota和越南仙草Mesona procumbens的净光合速率。李婷等[9]研究发现:光强过高或过低都会影响穿心莲Andrographis paniculata主要药用成分,随光强增加,穿心莲内酯(AND)量值变低,脱水穿心莲内酯(DDAND)变高。秦健等[10]发现青钱柳Cyclocarya paliurus幼苗随着光强增加其根茎叶生物量及总生物量呈现上升趋势。光照对三叶青的光合特性的影响研究已逐渐起步,刘崟艳等[11]通过不同产地三叶青植株的蒸腾作用与气孔结构研究,发现浙江台州等地的三叶青叶片均在弱光条件下时蒸腾速率最小、净光合速率最高;杨华等[12]研究了不同遮光条件下三叶青的光合特性,认为15%光照下其光合作用较好;钱丽华等[13]和彭昕等[14]发现光周期可调控三叶青叶片愈伤组织的生长及总黄酮量,结果显示12 h·d-1间断性光照最有利于愈伤组织的生长,24 h·d-1光照最有利于黄酮类化合物的累积。本研究着眼于研究不同光强下三叶青的光合指标的特性(净光合速率Pn,最大净光合速率Pmax,光补偿点LCP,光饱和点LSP,表观量子效率AQE,气孔导度Gs,胞间二氧化碳摩尔分数Ci,蒸腾速率Tr等),并进一步讨论其变化及规律,作为筛选三叶青生长的最佳光强条件依据,进行林下仿生栽培,有利于增加三叶青的产量,提高药材品质,满足医药需求。
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三叶青光合色素测定结果见表 1。经遮光处理后,三叶青叶绿素a,叶绿素b,总叶绿素和类胡萝卜素的量随着遮光程度的增加呈上升趋势。叶绿素a,叶绿素b,总叶绿素和类胡萝卜素的量在遮光50%以上显著增加,且在遮光90%出现最大值;叶绿素a/b值呈降低趋势,遮光50%以上差异显著(P < 0.05)。分析认为,光合色素的合成受光强影响,其中叶绿素在植物光合作用中具有吸收、传递和转化光能的作用,而类胡萝卜素为光吸收的辅助色素,对植物具有光保护作用。
表 1 不同光强处理对三叶青叶片色素的影响
Table 1. Effects of different light intensity on photosynthetic pigments content of Tetrastigma hemsleyanum with different light intensity treatments
遮光处理/% 三叶青叶片色素水平 w叶绿素a/(mg·g-1) w叶绿素b/(mg·g-1) w总叶绿素/(mg·g-1) w类胡萝卜素/(mg·g-1) 叶绿素a/b Ⅰ-ck 0.47 ± 0.05 bc 0.07 ± 0.01 c 0.54 ± 0.07 cd 0.15 ± 0.01 cd 7.06 ± 0.39 a Ⅰ-30 0.41 ± 0.04 b 0.06 ± 0.01 c 0.47 ± 0.05 d 0.13 ± 0.01 d 6.95 ± 0.54 a Ⅰ-50 0.54 ± 0.05 c 0.09 ± 0.01 b 0.63 ± 0.06 c 0.17 ± 0.04 c 5.98 ± 0.11 b Ⅰ-70 0.66 ± 0.05 d 0.12 ± 0.01 a 0.77 ± 0.06 b 0.20 ± 0.01 b 5.66 ± 0.28 b Ⅰ-90 0.81 ± 0.03 a 0.13 ± 0.01 a 0.94 ± 0.04 a 0.26 ± 0.005 a 6.27 ± 0.30 b Ⅱ-ck 0.41 ± 0.02 b 0.06 ± 0.02 0.47 ± 0.06 c 0.14 ± 0.02 c 7.49 ± 0.40 a Ⅱ-30 0.37 ± 0.04 b 0.05 ± 0.03 c 0.42 ± 0.04 c 0.12 ± 0.03 c 7.21 ± 0.50 a Ⅱ-50 0.60 ± 0.04 c 0.10 ± 0.03 b 0.70 ± 0.05 b 0.21 ± 0.03 b 5.86 ± 0.12 b Ⅱ-70 0.70 ± 0.04 d 0.13 ± 0.02 b 0.82 ± 0.06 a 0.21 ± 0.02 b 5.55 ± 0.30 b Ⅱ-90 0.84 ± 0.03 a 0.14 ± 0.03 a 0.98 ± 0.04 a 0.27 ± 0.01 a 6.15 ± 0.32 b 说明:不同字母表示在0.05水平上差异显著 在高温缓慢生长期,全光照(ck)和遮光30%的叶片色素量有所降低,叶绿素a/b比值增加,其余各组色素的量均上升,叶绿素a/b比值下降。叶绿素a的最大吸收在长波红光区(680 nm),叶绿素b的最大吸收在短波蓝光区(460 nm)。叶绿素a/b变小反映植物对外界短波光吸收量的相对增加, 说明在遮光30%以下三叶青叶片光合系统吸收过量的光能,导致光合色素的反应中心的失活或损害,遭受了强光胁迫,各生物量下降,合成光合色素需要的物质供给减少,因而光合色素总量降低,生长不适应;在遮光50%以上,未产生强光胁迫,生长较适应,也间接证明了三叶青具有较强的耐阴性。
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如图 1所示:各遮光处理三叶青的光合-光响应曲线变化趋势基本一致。总体而言,当PAR在0~200 μmol·m-2·s-1时,Pn迅速升高;PAR在200~400 μmol·m-2·s-1时,Pn上升速度减缓;当PAR > 600 μmol·m-2·s-1时,遮光50%以上的Pn趋于稳定,而遮光30%和ck的Pn呈现降低趋势。随遮光程度增强,Pn呈现先增后降趋势,遮光50%以上的Pn均高于ck,在遮光70%处理组达到最大值。处于高温缓慢生长期时,各处理下Pn均比快速生长期有所降低。
图 1 不同光强处理下三叶青光合-光响应曲线
Figure 1. Net photosynthetic rate (Pn)-light response curves of Tetrastigma hemsleyanum with different light intensity treatments
由表 2可以看出:LSP,AQE和Pmax随着遮光程度增加呈现先上升后降低趋势,均在遮光70%达到最大值;LCP呈现下降趋势,在遮光90%达到最小值,Rd的变化上下波动,但均在遮光70%时达到最大值。与快速生长期相比,高温缓慢生长期相同遮光处理下的LSP,AQE,Rd,Pmax数值均下降,而LCP则呈现上升趋势,与ck相比,LSP,AQE,Rd在50%以上差异显著(P < 0.05)。
表 2 不同光强处理下三叶青的光合响应曲线特征参数
Table 2. Characteristic parameters of photosynthetic response curve of Tetrastigma hemsleyanum with different light intensity treatments
遮光处理/% 光合参数 LSP/(μ·mol-2·s-1) LCP/(μ·mol-2·s-1) AQE/(mol·mol-1) Rd/(μ·mol-2·s-1) Pmax/(μ·mol-2·s-1) Ⅰ-ck 97.36 ± 19.23 c 12.67 ± 2.31 ab 0.070 ± 0.020 b 0.88 ± 0.03 b 2.88 ± 1.05 b Ⅰ-30 88.58 ± 14.68 c 14.65 ± 1.16 a 0.060 ± 0.010 b 0.87 ± 0.15 b 2.73 ± 0.73 b Ⅰ-50 224.48 ± 24.94 b 12.66 ± 3.05 ab 0.080 ± 0.030 b 0.82 ± 0.25 ab 3.53 ± 0.83 b Ⅰ-70 340.49 ± 32.11 a 8.66 ± 4.17 bc 0.180 ± 0.070 a 1.20 ± 0.41 b 5.34 ± 1.00 a Ⅰ-90 244.95 ± 33.03 b 6.99 ± 1.73 c 0.050 ± 0.020 b 0.42 ± 0.16 a 3.73 ± 0.89 ab Ⅱ-ck 76.71 ± 3.06 c 26.68 ± 10.27 a 0.030 ± 0.010 b 0.60 ± 0.05 b 1.19 ± 0.34 b Ⅱ-30 66.05 ± 0.01 c 18.68 ± 5.03 ab 0.030 ± 0.006 b 0.57 ± 0.08 b 1.21 ± 0.30 b Ⅱ-50 86.61 ± 23.87 bc 14.66 ± 4.16 bc 0.040 ± 0.002 ab 0.58 ± 0.21 b 1.88 ± 0.68 ab Ⅱ-70 159.28 ± 10.36 a 13.33 ± 4.61 bc 0.060 ± 0.006 a 0.70 ± 0.27 b 2.45 ± 0.71 a Ⅱ-90 105.97 ± 11.22 b 5.34 ± 4.17 c 0.041 ± 0.026 ab 0.17 ± 0.12 a 1.38 ± 0.54 b 说明:不同字母表示在0.05水平上差异显著 -
气孔是植物与外界进行气体交换和蒸腾作用的通道,因此,Gs的大小决定二氧化碳和水进出植物细胞的速率,进而间接影响植物的光合作用和蒸腾速率。由图 2可见:各处理均呈现先快速上升后趋于稳定的趋势,其中,遮光50%~90%处理的Gs均高于遮光30%和全光照,表明可能三叶青在长期的强光胁迫下可导致气孔关闭,故Gs下降。快速生长期的Gs最大值出现在遮光70%(54.33 mmol·m-2·s-1),高温缓慢生长期各处理下Gs在均有所降低,最大值为遮光50%(30.02 mmol·m-2·s-1)。分析认为,夏季的高温及高光强使三叶青生长滞缓,气孔开闭不活跃,各处理Gs均降低。
由图 3可知:当PAR在0~200 μmol·m-2·s-1时,不同遮光处理下三叶青的Ci快速下降,并在之后随光强增加,各处理的Ci处于低水平;不同生育时期的三叶青Ci差异不大。一般而言,Ci大小取决于4个因素:叶片周围空气中的二氧化碳浓度,Gs,叶肉导度(gm)和叶肉细胞的光合活性[18]。当叶片周围空气二氧化碳浓度升高,Gs和gm增大,叶肉细胞的光合活性降低时,Ci会增大;反之,Ci则减小。本研究中,不同遮光处理下三叶青的Ci先快速下降,后保持稳定,与光合响应曲线的变化趋势负相关。随着光照强度的增加,三叶青进行光合作用,二氧化碳快速消耗,因此,Ci降低,Pn快速增加;不同生育时期,三叶青叶片的Ci与Pn变化趋势一致,即都在高温缓慢生长期下降,显示该时期三叶青生长缓慢,Pn较低,对二氧化碳的消耗较少。
图 4显示:随着光照强度的增加,各处理的Tr均出现上升趋势。三叶青属于喜阴药用植物,遮光可减少高光强伤害,并使气孔活跃度增加[19],故Tr随Gs呈现上升趋势。在快速生长期,Tr最大值(0.98 mmol·m-2·s-1)出现在遮光70%处理组,高温缓慢生长期的Tr最大值(0.58 mmol·m-2·s-1)出现在遮光90%处理组,与Gs变化同步,或与盛夏高光强、高温及低空气湿度等因素有关[20]。
Effects of light intensity treatments on photosynthetic characteristics in Tetrastigma hemsleyanum
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摘要: 为探讨适宜三叶青Tetrastigma hemsleyanum生长的光强环境,并为其人工栽培和林下栽培提供基础,以2年生三叶青扦插苗为试验材料,对它们进行不同梯度的遮光处理,梯度设置为全光照(ck),遮光30%,50%,70%和90%,研究光强对三叶青不同生育时期(快速生长期与高温缓慢生长期)的光合特性的影响。采用体积分数为95%乙醇浸提法测定三叶青叶片中光合色素,利用LI-6400XT便携式光合系统测定仪测定叶片净光合速率(Pn),气孔导度(Gs),胞间二氧化碳摩尔分数(Ci)和蒸腾速率(Tr)等指标,计算光饱和点(LSP),光补偿点(LCP),量子效率(AQE),暗呼吸速率(Rd)和最大净光合速率(Pmax),并拟合光响应曲线。结果表明:随着遮光程度的增加,叶绿素a,叶绿素b,总叶绿素和类胡萝卜素总体上呈现上升趋势,但是在不同生育时期变化不大;多数光合指标亦随遮光程度增加而呈现增长趋势,多在遮光70%时达到最大值之后下降,且快速生长期(Pn为4.19 μmol·m-2·s-1,Gs为53.54 mol·m-2·s-1,Tr为0.98 mmol·m-2·s-1,LSP为340.49 μmol·m-2·s-1,AQE为0.18 mol·mol-1,Rd为1.20 μmol·m-2·s-1,Pmax为5.34 μmol·m-2·s-1)比高温缓慢生长期(Pn为1.74 μmol·m-2·s-1,Gs为17.53 mol·m-2·s-1,Tr为0.36 mmol·m-2·s-1,LSP为159.28 μmol·m-2·s-1,AQE为0.06 mol·mol-1,Rd为0.70 μmol·m-2·s-1,Pmax为2.45 μmol·m-2·s-1)数值高41.67%~67.26%;Ci和LSP呈现先下降后趋于稳定趋势,表明高温缓慢生长期的强光、高温、低湿等环境条件使其光合活跃度下降,对植株生长产生胁迫。综合试验结果得出结论,不同生育时期三叶青均在遮光度70%的条件下生长适宜。Abstract: Tetrastigma hemsleyanum has obvious pharmacological effects on inhibitting tumor cell proliferation and migration clinically. It is on the verge of extinction with the increasing demand over the past decade, so bionic planting is imperative to meet the medical needs. In this paper, we explored suitable light intensity conditions for the T. hemsleyanum growth by consideration that T. hemsleyanum grows in the forest wildly and is sensitive to strong light. Two-year-old T. hemsleyanum cuttings were shaded with different gradients treatment as test materials. The shading gradients were set as full light(ck), shade 30%, 50%, 70% and 90%. The responses of photosynthetic characteristics to light intensity were analyzed in different growth stages (rapid growth period and slow growth period). Following indicators were also measured:contents of photosynthetic pigments were determined by 95% alcohol extraction; light photosynthetic rate (Pn), stomatal conductance (Gs), intercellular CO2 concentration (Ci) and transpiration rate (Tr) were measured by LI-6400XT portable photosynthesis system; light response curve (LSP), light compensation point (LSP), quantum efficiency (AQE), dark respiration rate (Rd) and maximum net photosynthetic rate (Pmax) were calculated, and the light response curve was fitted. Results showed that trends of chlorophyll a (Chla), chlorophyll b (Chlb), total chlorophyll (Chl) and carotenoid (Car) were increased with the increase of shade gradients, but the contents of photosynthetic pigment changed little in different periods; most photosynthetic indices also showed increasing trends with the increase of shading gradients, reached the maximum values in the treatment of 70% shade, then reduced, the values of rapid growth period (Pn was 4.19 μmol·m-2·s-1, Gs was 53.54 mol·m-2·s-1, Tr was 0.98 mmol·m-2·s-1, LSP was 340.49 μmol·m-2·s-1, AQE was 0.18 mol·mol-1, Rd was 1.20 μmol·m-2·s-1 and Pmax was 5.34 μmol·m-2·s-1) were 41.67%~67.26% higher than those of slow growth period (Pn was 1.74 μmol·m-2·s-1, Gs was 17.53 mol·m-2·s-1, Tr was 0.36 mmol·m-2·s-1, LSP was 159.28 μmol·m-2·s-1, AQE was 0.06 mol·mol-1, Rd was 0.70 μmol·m-2·s-1 and Pmax was 2.45 μmol·m-2·s-1); Ci and LSP showed trends of decreasing first and then tending to be stable. It showed that the photosynthetic activity decreased and plant growth slowed down with unsuitable environmental conditions such as strong light, high temperature and low humidity. The analysis of results indicated that the growth of 70% shading was appropriate in different growth periods. This paper discussed the relationship between physiological characteristics and light intensity of T. hemsleyanum in different growth periods, enriches the research results of physiological aspects, and provided basic theory for artificial cultivations and under-forest-cultivation.
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Key words:
- botany /
- Tetrastigma hemsleyanum /
- shade /
- photosynthetic pigments /
- photosynthetic characteristics
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表 1 不同光强处理对三叶青叶片色素的影响
Table 1. Effects of different light intensity on photosynthetic pigments content of Tetrastigma hemsleyanum with different light intensity treatments
遮光处理/% 三叶青叶片色素水平 w叶绿素a/(mg·g-1) w叶绿素b/(mg·g-1) w总叶绿素/(mg·g-1) w类胡萝卜素/(mg·g-1) 叶绿素a/b Ⅰ-ck 0.47 ± 0.05 bc 0.07 ± 0.01 c 0.54 ± 0.07 cd 0.15 ± 0.01 cd 7.06 ± 0.39 a Ⅰ-30 0.41 ± 0.04 b 0.06 ± 0.01 c 0.47 ± 0.05 d 0.13 ± 0.01 d 6.95 ± 0.54 a Ⅰ-50 0.54 ± 0.05 c 0.09 ± 0.01 b 0.63 ± 0.06 c 0.17 ± 0.04 c 5.98 ± 0.11 b Ⅰ-70 0.66 ± 0.05 d 0.12 ± 0.01 a 0.77 ± 0.06 b 0.20 ± 0.01 b 5.66 ± 0.28 b Ⅰ-90 0.81 ± 0.03 a 0.13 ± 0.01 a 0.94 ± 0.04 a 0.26 ± 0.005 a 6.27 ± 0.30 b Ⅱ-ck 0.41 ± 0.02 b 0.06 ± 0.02 0.47 ± 0.06 c 0.14 ± 0.02 c 7.49 ± 0.40 a Ⅱ-30 0.37 ± 0.04 b 0.05 ± 0.03 c 0.42 ± 0.04 c 0.12 ± 0.03 c 7.21 ± 0.50 a Ⅱ-50 0.60 ± 0.04 c 0.10 ± 0.03 b 0.70 ± 0.05 b 0.21 ± 0.03 b 5.86 ± 0.12 b Ⅱ-70 0.70 ± 0.04 d 0.13 ± 0.02 b 0.82 ± 0.06 a 0.21 ± 0.02 b 5.55 ± 0.30 b Ⅱ-90 0.84 ± 0.03 a 0.14 ± 0.03 a 0.98 ± 0.04 a 0.27 ± 0.01 a 6.15 ± 0.32 b 说明:不同字母表示在0.05水平上差异显著 表 2 不同光强处理下三叶青的光合响应曲线特征参数
Table 2. Characteristic parameters of photosynthetic response curve of Tetrastigma hemsleyanum with different light intensity treatments
遮光处理/% 光合参数 LSP/(μ·mol-2·s-1) LCP/(μ·mol-2·s-1) AQE/(mol·mol-1) Rd/(μ·mol-2·s-1) Pmax/(μ·mol-2·s-1) Ⅰ-ck 97.36 ± 19.23 c 12.67 ± 2.31 ab 0.070 ± 0.020 b 0.88 ± 0.03 b 2.88 ± 1.05 b Ⅰ-30 88.58 ± 14.68 c 14.65 ± 1.16 a 0.060 ± 0.010 b 0.87 ± 0.15 b 2.73 ± 0.73 b Ⅰ-50 224.48 ± 24.94 b 12.66 ± 3.05 ab 0.080 ± 0.030 b 0.82 ± 0.25 ab 3.53 ± 0.83 b Ⅰ-70 340.49 ± 32.11 a 8.66 ± 4.17 bc 0.180 ± 0.070 a 1.20 ± 0.41 b 5.34 ± 1.00 a Ⅰ-90 244.95 ± 33.03 b 6.99 ± 1.73 c 0.050 ± 0.020 b 0.42 ± 0.16 a 3.73 ± 0.89 ab Ⅱ-ck 76.71 ± 3.06 c 26.68 ± 10.27 a 0.030 ± 0.010 b 0.60 ± 0.05 b 1.19 ± 0.34 b Ⅱ-30 66.05 ± 0.01 c 18.68 ± 5.03 ab 0.030 ± 0.006 b 0.57 ± 0.08 b 1.21 ± 0.30 b Ⅱ-50 86.61 ± 23.87 bc 14.66 ± 4.16 bc 0.040 ± 0.002 ab 0.58 ± 0.21 b 1.88 ± 0.68 ab Ⅱ-70 159.28 ± 10.36 a 13.33 ± 4.61 bc 0.060 ± 0.006 a 0.70 ± 0.27 b 2.45 ± 0.71 a Ⅱ-90 105.97 ± 11.22 b 5.34 ± 4.17 c 0.041 ± 0.026 ab 0.17 ± 0.12 a 1.38 ± 0.54 b 说明:不同字母表示在0.05水平上差异显著 -
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