Anatomical structure and physiological characteristics of ivy with formaldehyde and weak light stress

ZHANG Yongfu; MO Liling; NIU Yanfen; XIA Tiyuan; WANG Dingkang; GENG Kaiyou

doi:10.11833/j.issn.2095-0756.2016.06.013

Volume 33 Issue 6

Nov. 2016

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ZHANG Yongfu, MO Liling, NIU Yanfen, XIA Tiyuan, WANG Dingkang, GENG Kaiyou. Anatomical structure and physiological characteristics of ivy with formaldehyde and weak light stress[J]. Journal of Zhejiang A&F University, 2016, 33(6): 1017-1024. doi: 10.11833/j.issn.2095-0756.2016.06.013

Citation:

ZHANG Yongfu, MO Liling, NIU Yanfen, XIA Tiyuan, WANG Dingkang, GENG Kaiyou. Anatomical structure and physiological characteristics of ivy with formaldehyde and weak light stress[J]. Journal of Zhejiang A&F University, 2016, 33(6): 1017-1024. doi: 10.11833/j.issn.2095-0756.2016.06.013

Anatomical structure and physiological characteristics of ivy with formaldehyde and weak light stress

doi: 10.11833/j.issn.2095-0756.2016.06.013

1.
School of Agriculture, Kunming University, Kunming 650214, Yunnan, China
2.
Office of Science Research, Kunming University, Kunming 650214, Yunnan, China

Received Date: 2015-12-17
Rev Recd Date: 2016-01-25
Publish Date: 2016-12-20

Abstract

To determine the effects of formaldehyde and weak light stress on anatomical and physiological characteristics of two species of ivy(Hedera helix and Hedera nepalensis) with their adaptability, potted seedlings were used as experimental materials. Sealed glass containers held three stress treatments of formaldehyde, weak light, and formaldehyde+weak light, and a control. After four months of treatment, the anatomical structure of leaves, the physiological and biochemical indexes of roots and leaves were measured. Results for H. helix showed that with formaldehyde and with weak light stress, palisade tissue thickness, length and width of stomata decreased significantly (P<0.05); whereas spongy tissue thickness increased with formaldehyde stress, and spongy tissue density increased with weak light. However, for H. nepalensis and stress, thickness of the leaf, palisade tissue, and spongy tissue; and the length, width, and density of stomata all decreased. For both ivies, root vigor with all stresses decreased significantly (P<0.05). With formaldehyde and formaldehyde+weak light stresses, formaldehyde content of H. helix was higher than H. nepalensis. Also, with formaldehyde stress, chlorophyll content in H. nepalensis decreased, but it increased with weak light which was contrary to H. helix. In addition, with the formaldehyde+weak light stress treatment, chlorophyll content in both ivies decreased significantly (P<0.05), but carotenoid content increased significantly (P<0.05). In both ivies with the weak light treatment, superoxide dismutase (SOD) activity of roots and leaves decreased, and with formaldehyde and double stress the SOD activity in roots and leaves along with peroxidase (POD) activity in roots decreased significantly (P<0.05). Finally, for both ivies with all stresses, hydrogen peroxide, malondialdehyde (MDA), relative electrical conductivity, and the oxygen free radical generation rate increased. Thus, the adaptability of H. nepalensis to weak light was stronger than H. helix, but the resistance to formaldehyde was contary.[Ch, 5 fig. 1 tab. 21 ref.]
- botany,
- Hedera spp.,
- formaldehyde stress,
- weak light stress,
- anatomic structure,
- physiological response

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References

[1]	BAZZAZ F A. Plants in Changing Environments:Linking Physiological,Population, and Community Ecology[M]. Cambridge:Cambridge University Press, 1996.
[2]	CALLAHAN H S, PIGLIUCCI M. Shade-induced plasticity and its ecological significance in wild populations of Arabidopsis thaliana[J]. Ecology, 2002, 83(7):1965-1980.
[3]	CHEN Xiong, WU Dongxiu, WANG Genxuan, et al. Effect of elevated CO₂ concentration on photosynthesis and antioxidative enzyme activities of wheat plant grown under drought condition[J]. Chin J Appl Ecol, 2000, 11(6):881-884.
[4]	HE Weiming, ZHONG Zhangcheng. Morphological and growth responses of the climbing plant, Gynostemma pentaphllum seedlings to varying light intensity[J]. Acta Phytoecol Sin, 2000, 24(3):375-378.

[6]	LIU Yanli, CHEN Nengchang, ZHOU Jianmin, et al. Researches in the removal of formaldehyde from indoor air by ornamental plant[J]. Ind Catalys, 2008, 16(9):6-11.
[7]	CAO Yonghui, ZHOU Benzhi, CHEN Shuanglin. Effects of water stress on physiological characteristics of different illicium lanceolatum ecotypes under low light intensity[J]. Acta Ecol Sin, 2014, 34(4):814-822.
[8]	MENG Yaning,YAN Libin,FAN Yanqin. Research on changes of three enzymes activities of sweet(hot) pepper under low temperature and poor light and the tolerance[J]. J Hebei Agric Sci, 2014, 18(6):21-24, 85.
[9]	XUAN Xiuxia, XIAO Suqin, YOU Lan,et al. The analysis of the intermediates for formaldehyde metabolism and physiological changes under gaseous formaldehyde stress in Hedera helix[J]. Life Sci Res, 2013, 17(2):125-135.
[10]	LINGHU Yuwei, LI Bin, LI Sifeng, et al. Monitoring, purification and response of three indoor ornamental plants on formaldehyde pollution[J]. Acta Bot Boreal-Occident Sin, 2011, 31(4):776-782.
[11]	ZHANG Shulin. Study on the methods for determining trace formaldehyde by phloroglucinol[J]. Environ Eng, 2005, 23(4):77-79.


[14]	MACDONALD S E, CHINNAPPA C C, REID D M. Evolution of phenotypic plasticity in the Stellaria longipes complex:Comparisons among cytotypes and habitats[J]. Evolution, 1988, 42(5):1036-1046.
[15]	DI Bao, CHEN Duanfen, GUO Xiumin, et al. Effect of leaf structure on the formaldehyde-resistance capacity in five ornamental plants[J]. Northern Hortic, 2007(10):122-124.
[16]	XIAO Jian, WU Sizheng, NIE Dongling, et al. Effects of formaldehyde stress on physiology and biochemistry of three ornamental plants[J]. J Hunan For Sci Technol, 2012, 39(2):45-48.
[17]	GUO Jinping, LI Haibo, LIU Ning, et al. Comparison of responses of Larix principis-rupprechtii and Picea meyeri seedling growth to light availability and planting density under controlled environment[J]. Sci Silv Sin, 2009, 45(2):53-59.
[18]	LI Chaohai, LUAN Limin, YIN Fei, et al. Effects of light stress at different stages on the growth and yield of different maize genotypes (Zea mays L.)[J]. Acta Ecol Sin, 2005, 25(4):824-830.
[19]	CHAO Tiancai, ZHOU Shoubiao, CHANG Linlin, et al. Effects of light intensity on the leaf morphology and physiological parameters of Boea clarkeana[J]. Chin J Ecol, 2013, 32(5):1161-1167.
[20]	CHU Ling, LIU Dengyi, WANG Youbao, et al. Separate and combined effects of Cu and Cd on seedling growth and active oxygen metabolism system of Trifolium repens L.[J]. Front Biosci, 2006, 11(11):2861-2867.
[21]	LIU Dong, SHI Baosheng, WEI Wenxin. Effect of formaldehyde gas stress on morphology and partial physiological indexes of three ornamental plants[J]. J Agric Univ Hebei, 2011, 34(2):66-70.

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沈阳化工大学材料科学与工程学院沈阳 110142

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光作为重要的环境因子，对植物生长、发育和演化具有极其重要的作用，同时又影响植物形态和生理功能^[1]。光照强度长时间大幅度低于植物光饱和点将产生弱光胁迫，影响植物的形态建成和生理生化过程^[2-3]。弱光影响植物最直接的生理表现是光合速率降低^[4]，同时植物激素、抗氧化剂及次生代谢等生理进程也受到明显影响。甲醛是一种无色、有强烈刺激性气味的气体，为挥发性有机物，易溶于水，在室温下易释放。世界卫生组织指出，当室内甲醛超过0.1 mg·m^-3时，就应当引起人们足够的重视^[5]，许多建筑材料和胶黏剂均含有甲醛，室内装修后甲醛会不断向环境释放，释放周期可长达10余年。室内装修材料及生活用品带来的甲醛严重威胁着人们的身体健康而广受关注^[6]。目前，可通过物理、化学、生物工程以及绿色植物净化方法控制室内甲醛污染，由于前3种方法效果不理想且局限性较大，因此利用植物吸收室内装修产生的甲醛成为当今研究的热点。室内光照强度明显低于室外，植物的生长发育势必受到较大的影响。国内外相关研究大多局限于弱光^[7-8]或甲醛^[9-10]等单一因子对植株生长发育和生理生态等方面的影响，有关甲醛和光照强度交互作用对植物解剖结构和生理生态特性及相关适应性影响的研究还未见报道。常春藤 Hedera是常见的常绿木质藤本植物，对环境的适应性强，适于室内外栽植，叶有香气，形态优美，且易繁殖，已成为主要的观赏植物之一。目前，对常春藤甲醛吸收能力已有广泛的研究，但对弱光下常春藤对甲醛胁迫的解剖特征及生理响应的研究还未见报道。因此，本研究选取中华常春藤Hedera nepalensis和洋常春藤Hedera helix半年生扦插盆栽苗进行甲醛和弱光环境培养试验，探讨2种常春藤苗抗性系统对一定范围弱光环境和甲醛胁迫影响的解剖及生理响应，揭示它们对甲醛和弱光胁迫的适应和调节机制，以及甲醛胁迫在植物忍耐弱光适应性调节过程中发挥的作用，以期为常春藤室内栽培和光照提供理论依据。

1. 材料与方法

1.1. 材料培养

供试材料为昆明学院校园内边坡地被植物中华常春藤Hedera nepalensis（简写为HN）和洋常春藤H. helix（简写为HH）的半年生扦插苗。选取根系健壮，枝叶繁茂的植株移栽至20 cm × 20 cm的营养钵中，然后转移到100 cm × 60 cm × 100 cm的玻璃容器中，各个玻璃容器分别放置中华常春藤和洋常春藤各10盆，且分别浇Hoagland营养液1次200 mL ·盆^-1·周^-1 （pH 5.5），以维持植株正常生长。

1.2. 试验设计

植物材料在玻璃容器中正常培养2周后，分别进行以下4个处理：T₁为甲醛处理，T₂弱光处理，T₃为甲醛 + 弱光处理，ck为对照，重复3次·处理^-1。T₁和T₃甲醛处理的方法为在玻璃容器中央悬挂1个脱脂棉球，把1.0 mL 体积分数为40%的甲醛溶液滴到脱脂棉球上，甲醛不能下滴，让其挥发，然后玻璃容器的上方用玻璃板和密封圈封好，防止容器内的甲醛挥发出来。T₂和T₃用遮阳网罩住玻璃容器进行遮光处理，遮光的光照强度是未遮光的1/10。处理4个月后，采样分别测定叶的解剖结构及根、叶的生理生化指标。甲醛的量和弱光的光照强度由预实验确定。

1.3. 测定指标及方法

叶片横切结构观察的方法：取成熟叶片用剪刀沿中脉2侧剪成0.5 cm × 0.5 cm的小块，置于标准固定液（FAA）中固定48 h，梯度乙醇逐级脱水，苯胺番红和苯胺固绿对染，二甲苯透明，常规石蜡切片，厚度为8 μm，在Olympus光学显微镜下观察、摄影及测量。

叶表皮气孔观察的方法：上述固定后的叶片小块，置于铬酸-硝酸离析液中离析24 h，用清水洗净，番红染液染色，梯度乙醇脱水，二甲苯透明后制片，在Olympus光学显微镜观察、摄影及测量。

甲醛质量分数的测定参照张书林的方法^[11]；根系活力的测定采用氯化三苯基四氮唑（TTC）-琥珀酸钠显色法，叶绿素和类胡萝卜素质量分数的测定采用体积分数为96%乙醇提取法，超氧化物歧化酶（SOD）活性采用氮蓝四唑（NBT）光反应法测定，过氧化氢酶活性采用硫酸钛-浓氨水法测定，相对电导率采用电导仪测定法，丙二醛质量摩尔浓度采用硫代巴比妥酸法测定^[12]。此外，过氧化物酶活性的测定采用愈创木酚-双氧水显色法，氧自由基产生速率的测定采用对氨基苯磺酸-α-萘胺显色法^[13]。

1.4. 数据统计分析

所有数据用统计软件SPSS 19.0 在0.05 水平上进行单因素方差分析，用Excel 2007制图。

3. 结论与讨论

植物表型可塑性是由于环境变化引起基因表达的变化从而产生不同的表型，是植物对环境变化的重要适应^[14]。植物叶器官受环境影响较大，通过对中华常春藤和洋常春藤进行长达4个月的甲醛和弱光处理培养，2种常春藤的叶片解剖结构均发生了较大的变化，主要表现在甲醛和弱光处理使叶片厚度、栅栏组织厚度减小，其中弱光处理的减小幅度最大。气孔是植物与环境之间气体交换的通道，空气中的各种污染物主要通过气孔进入植物组织，气孔在叶肉组织与大气之间构成了一个可变的扩散阻力，调控气体的吸收，气孔开度缩小或关闭可增强植物对大气污染的抗性^[15]。从本研究结果来看，甲醛和弱光处理均使常春藤的气孔长度、宽度和密度降低，其中弱光和双重处理下降达到差异显著（P＜0.05）性水平；中华常春藤的气孔长度、宽度和密度在各处理下均大于洋常春藤，但结合植物体内的甲醛质量分数来看，甲醛处理后洋常春藤根系和叶片的甲醛质量分数高于中华常春藤。可见，尽管在甲醛和弱光下，常春藤叶片解剖结构发生了一系列的变化响应，但对逆境的抵抗能力更多还是通过生理生化调控来实现的。

甲醛和弱光胁迫均是影响植株生长发育和光合作用的逆境因素。甲醛胁迫可使植物的叶绿素质量分数降低^[16]，原因是甲醛胁迫下，抗氧化酶系统清除氧自由基能力降低，引起细胞膜脂过氧化，丙二醛不断积累，从而破坏了叶绿体的膜系统，叶绿素降解且质量分数大幅降低^[10]。从高光环境到低光环境，植物所能获得的光量减少。为了更有效地捕获光能，植物会相应地增加叶内的叶绿素质量分数^[17]。生长在弱光环境下的植物，叶绿素质量分数增加使其能够充分吸收光合有效辐射^[18]，但光照太弱也不利于叶绿素的合成。从本研究来看，甲醛胁迫下中华常春藤的叶绿素a，叶绿素b和叶绿素a+b质量分数均下降，而洋常春藤则明显上升，说明甲醛对洋常春藤叶绿素合成有促进作用；另一方面，在弱光下中华常春藤的叶绿素a，叶绿素b和叶绿素a+b质量分数均上升，而洋常春藤则显著（P＜0.05）下降，说明中华常春藤对弱光的适应性更强；此外，在甲醛和弱光双重胁迫下，2种常春藤的叶绿素a，叶绿素b和叶绿素a+b质量分数均显著（P＜0.05）下降，类胡萝卜素质量分数则显著（P＜0.05）上升，说明双重胁迫对2种常春藤均产生了较大的影响。

通过CAT，POD 和SOD活性的动态变化研究发现，弱光下CAT，POD 及SOD 活性相对处于较低的状态，说明当活性氧产生速率较低时，保护酶活性也处于一个相对较低的状态^[19]。植物体内积累了过量活性氧自由基可引发膜脂过氧化，MDA是膜脂过氧化的主要产物之一，其大量积累是活性氧伤害作用的表现。细胞膜通透性的大小可作为植物对不良环境胁迫的一个指标^[20]。研究发现，弱光下2种常春藤根系和叶片中的SOD活性下降，而根系中的 POD活性也下降，叶片中则上升；弱光下根系的过氧化氢质量摩尔浓度、氧自由基产生速率和相对电导率显著（P＜0.05）上升，而叶片差异不大；此外，弱光还显著（P＜0.05）增加了根系和叶片的MDA质量分数。甲醛胁迫可破坏植物细胞膜，使其透性增大，渗透物质外流。本研究中2种常春藤的细胞膜透性在甲醛胁迫下均增大，这与刘栋等^[21]的研究结果相同。在甲醛胁迫下，2种常春藤根系和叶片的SOD活性、过氧化氢质量摩尔浓度、氧自由基产生速率和MDA质量分数上升，POD活性显著（P＜0.05）降低，这与令狐昱慰等^[10]的研究结果基本一致。同时，本研究结果还发现，甲醛和弱光双重胁迫使2种常春藤根系和叶片的SOD活性、氧自由基产生速率、过氧化氢和MDA质量分数上升，而POD活性显著（P＜0.05）降低。

从本研究结果来看，中华常春藤对弱光的适应性强于洋常春藤，而洋常春藤对甲醛的抗性强于中华常春藤，且洋常春藤单位组织吸收的甲醛量大于中华常春藤。此外，甲醛和弱光双重胁迫下虽然对2种常春藤的生长生理均造成较大的伤害，但2种常春藤在叶片解剖和生理生化方面做出积极响应去适应这些逆境胁迫。

Reference (21)

[1]	BAZZAZ F A. Plants in Changing Environments:Linking Physiological,Population, and Community Ecology[M]. Cambridge:Cambridge University Press, 1996.
[2]	CALLAHAN H S, PIGLIUCCI M. Shade-induced plasticity and its ecological significance in wild populations of Arabidopsis thaliana[J]. Ecology, 2002, 83(7): 1965-1980. doi: 10.1890/0012-9658(2002)083[1965:SIPAIE]2.0.CO;2
[3]	陈雄, 吴冬秀, 王根轩. CO₂浓度升高对干旱胁迫下小麦光合作用和抗氧化酶活性的影响[J]. 应用生态学报, 2000, 11(6): 881-884.	CHEN Xiong, WU Dongxiu, WANG Genxuan. Effect of elevated CO₂ concentration on photosynthesis and antioxidative enzyme activities of wheat plant grown under drought condition[J]. Chin J Appl Ecol, 2000, 11(6): 881-884.
[4]	何维明, 钟章成. 攀援植物绞股蓝幼苗对光照强度的形态和生长反应[J]. 植物生态学报, 2000, 24(3): 375-378.	HE Weiming, ZHONG Zhangcheng. Morphological and growth responses of the climbing plant, Gynostemma pentaphllum seedlings to varying light intensity[J]. Acta Phytoecol Sin, 2000, 24(3): 375-378.
[5]	朱天乐. 室内空气污染控制[M]. 北京:化学工业出版社, 2003:46-47.
[6]	刘艳丽, 陈能场, 周建民. 观赏植物净化室内空气中甲醛的研究进展[J]. 工业催化, 2008, 16(9): 6-11.	LIU Yanli, CHEN Nengchang, ZHOU Jianmin. Researches in the removal of formaldehyde from indoor air by ornamental plant[J]. Ind Catalys, 2008, 16(9): 6-11.
[7]	曹永慧, 周本智, 陈双林. 弱光下水分胁迫对不同产地披针叶茴香幼苗生理特性的影响[J]. 生态学报, 2014, 34(4): 814-822.	CAO Yonghui, ZHOU Benzhi, CHEN Shuanglin. Effects of water stress on physiological characteristics of different illicium lanceolatum ecotypes under low light intensity[J]. Acta Ecol Sin, 2014, 34(4): 814-822.
[8]	孟雅宁, 严立斌, 范妍芹. 低温弱光下甜(辣)椒3种酶活性的变化及其耐受性研究[J]. 河北农业科学, 2014, 18(6): 21-24.	MENG Yaning, YAN Libin, FAN Yanqin. Research on changes of three enzymes activities of sweet(hot) pepper under low temperature and poor light and the tolerance[J]. J Hebei Agric Sci, 2014, 18(6): 21-24.
[9]	轩秀霞, 肖素勤, 游览. 常春藤代谢气体甲醛中间产物及甲醛胁迫下叶片相关生理特性变化分析[J]. 生命科学研究, 2013, 17(2): 125-135.	XUAN Xiuxia, XIAO Suqin, YOU Lan. The analysis of the intermediates for formaldehyde metabolism and physiological changes under gaseous formaldehyde stress in Hedera helix[J]. Life Sci Res, 2013, 17(2): 125-135.
[10]	令狐昱慰, 黎斌, 李思锋. 3种观赏植物对室内甲醛污染的净化及生长生理响应[J]. 西北植物学报, 2011, 31(4): 776-782.	LINGHU Yuwei, LI Bin, LI Sifeng. Monitoring, purification and response of three indoor ornamental plants on formaldehyde pollution[J]. Acta Bot Boreal-Occident Sin, 2011, 31(4): 776-782.
[11]	张书林. 间苯三酚比色法测定微量甲醛研究[J]. 环境工程, 2005, 23(4): 77-79.	ZHANG Shulin. Study on the methods for determining trace formaldehyde by phloroglucinol[J]. Environ Eng, 2005, 23(4): 77-79.
[12]	邹琦. 植物生理学实验指导[M]. 北京:中国农业出版社, 2000.
[13]	李合生. 植物生理生化实验原理和技术[M]. 北京:高等教育出版社, 2000.
[14]	MACDONALD S E, CHINNAPPA C C, REID D M. Evolution of phenotypic plasticity in the Stellaria longipes complex:Comparisons among cytotypes and habitats[J]. Evolution, 1988, 42(5): 1036-1046. doi: 10.2307/2408919
[15]	邸葆, 陈段芬, 果秀敏. 几种花卉叶片的组织结构对其抗甲醛能力的影响[J]. 北方园艺, 2007, (10): 122-124.	DI Bao, CHEN Duanfen, GUO Xiumin. Effect of leaf structure on the formaldehyde-resistance capacity in five ornamental plants[J]. Northern Hortic, 2007, (10): 122-124.
[16]	肖健, 吴思政, 聂东伶. 甲醛胁迫对3种室内观赏植物的生理生化影响[J]. 湖南林业科技, 2012, 39(2): 45-48.	XIAO Jian, WU Sizheng, NIE Dongling. Effects of formaldehyde stress on physiology and biochemistry of three ornamental plants[J]. J Hunan For Sci Technol, 2012, 39(2): 45-48.
[17]	郭晋平, 李海波, 刘宁. 华北落叶松和白杄幼苗对光照和竞争响应的差异比较[J]. 林业科学, 2009, 45(2): 53-59.	GUO Jinping, LI Haibo, LIU Ning. Comparison of responses of Larix principis-rupprechtii and Picea meyeri seedling growth to light availability and planting density under controlled environment[J]. Sci Silv Sin, 2009, 45(2): 53-59.
[18]	李潮海, 栾丽敏, 尹飞. 弱光胁迫对不同基因型玉米生长发育和产量的影响[J]. 生态学报, 2005, 24(5): 824-830.	LI Chaohai, LUAN Limin, YIN Fei. Effects of light stress at different stages on the growth and yield of different maize genotypes (Zea mays L.)[J]. Acta Ecol Sin, 2005, 24(5): 824-830.
[19]	晁天彩, 周守标, 常琳琳. 光照强度对大花旋蒴苣苔叶形态和生理指标的影响[J]. 生态学杂志, 2013, 32(5): 1161-1167.	CHAO Tiancai, ZHOU Shoubiao, CHANG Linlin. Effects of light intensity on the leaf morphology and physiological parameters of Boea clarkeana[J]. Chin J Ecol, 2013, 32(5): 1161-1167.
[20]	CHU Ling, LIU Dengyi, WANG Youbao. Separate and combined effects of Cu and Cd on seedling growth and active oxygen metabolism system of Trifolium repens L[J]. Front Biosci, 2006, 11(11): 2861-2867.
[21]	刘栋, 史宝胜, 魏文欣. 甲醛气体胁迫对3种观赏植物的形态及部分生理指标的影响[J]. 河北农业大学学报, 2011, 34(2): 66-70.	LIU Dong, SHI Baosheng, WEI Wenxin. Effect of formaldehyde gas stress on morphology and partial physiological indexes of three ornamental plants[J]. J Agric Univ Hebei, 2011, 34(2): 66-70.

处理	叶片厚度/μm	栅栏组织厚度/μm	海绵组织厚度/μm	气孔长度/μm	气孔宽度/μm	气孔密度/(个·mm^-2)
T₁-HN	316.37±27.33 b	133.08±11.21b	151.18±17.23ab	24.64±2.44ab	7.03±0.74ab	321.12±34.31a
T₁-HH	268.68±41.09c	112.46±9.87 c	129.67±15.01b	21.06±3.01b	6.55±1.23b	226.33±98.87 bc
T₂-HN	236.37±19.08c	85.27±10.06e	124.85±16.20b	18.64±1.55b	5.89±0.37bc	285.19±28.76 b
T₂-HH	179.09±9.76d	68.90±7.19e	84.22±9.06 c	16.32±1.38c	5.67±0.46 bc	301.56±42.07 a
T₃-HN	259.22±30.17c	80.81±10.33e	150.75±17.16ab	16.89±2.11c	5.37±0.85 c	215.07±28.16bc
T₃-HH	218.87±21.06c	71.57±8.72de	119.28±9.65 b	12.64±0.89d	4.43±1.08c	204.42±37.11c
ck-HN	398.48±33.71a	187.01±20.83 a	179.27±20.87 a	35.24±4.86 a	8.23±1.34a	345.86±54.21a
ck-HH	243.90±21.35c	119.12±13.27bc	96.58±11.07c	28.85±4.78 a	7.89±0.81 ab	275.31±38.09 b
说明：表中数据后的不同字母表示处理间差异达0.05显著水平。

Anatomical structure and physiological characteristics of ivy with formaldehyde and weak light stress

doi: 10.11833/j.issn.2095-0756.2016.06.013