不同砧木对‘87-1’葡萄光合特性及荧光特性的影响

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韩晓, 王海波, 王孝娣, 冀晓昊, 史祥宾, 王宝亮, 郑晓翠, 王志强, 刘凤之. 不同砧木对‘87-1’葡萄光合特性及荧光特性的影响[J]. 中国农业科学, 2018, 51(10): 1972-1981 https://doi.org/10.3864/j.issn.0578-1752.2018.10.016
HAN Xiao, WANG HaiBo, WANG XiaoDi, JI XiaoHao, SHI XiangBin, WANG BaoLiang, ZHENG XiaoCui, WANG ZhiQiang, LIU FengZhi. Effects of Different Rootstocks on ‘87-1’ Grape Photosynthetic and Chlorophyll Fluorescence Characteristics[J]. Scientia Acricultura Sinica, 2018, 51(10): 1972-1981 https://doi.org/10.3864/j.issn.0578-1752.2018.10.016

0 引言

【研究意义】光合作用是果树生长和结果的基础,果树的光合性能不仅受品种遗传性的制约,而且还受砧木特性的影响^[1]。‘87-1’葡萄作为设施栽培良种,在设施内已经大量种植。但是绝大多数都以‘贝达’做砧木,1103P、5C、华葡1号等优良砧木对‘87-1’光合特性和荧光特性的影响尚不明确。研究不同砧木对‘87-1’光合特性和荧光特性的影响,有助于筛选适宜设施栽培的优良砧穗组合。【前人研究进展】葡萄砧木研究始于葡萄根瘤蚜之后,葡萄根部害虫根瘤蚜从美洲传入欧洲并迅速传播,使欧洲葡萄园遭受近乎毁灭性的打击。后来研究证明美洲种的野生葡萄抗根瘤蚜,此后,对美洲葡萄野生种葡萄进行了改良,培育新的砧木品种,从而挽救了欧洲葡萄。葡萄砧木的应用挽救了世界上主要产区的葡萄生产,从此,对砧木的研究利用相继展开。国内外****发现不同砧木对葡萄成活率、物候期、果实品质、抗性、光合能力等^{[1,2,3,4,5,6,7,8,9,10]}方面均有显著影响,关于砧木对接穗光合特性的影响,主要集中在露地主栽品种的净光合速率、气孔导度、蒸腾速率等方面^{[11,12,13,14]}。【本研究切入点】由于用不同砧木嫁接适宜设施栽培葡萄品种的研究刚刚开始,生产者需要了解不同砧木的特性及其对嫁接品种的影响,以便选择适宜设施环境的砧木,而中国在这方面的研究几近空白;国外对设施葡萄砧木的评价筛选也尚未见报道。【拟解决的关键问题】本研究通过光补偿点、CO₂补偿点、暗呼吸速率、不同温度下净光合速率及叶绿素荧光等可以反应设施光合特性的参数,研究不同砧木对接穗耐弱光能力、耐低浓度CO₂能力、耐高温能力、叶绿素荧光特性的影响;综合评价8种常用砧木对‘87-1’光合特性和荧光特性的影响,初步为筛选适宜设施栽培的‘87-1’砧穗组合提供理论基础。

1 材料与方法

1.1 试验材料及地点

试验于2016—2017年连续两年的6—7月（花期和幼果发育期）在中国农业科学院果树研究所葡萄核心技术试验示范园（东经120.51°,北纬40.45°）进行。示范园土壤类型为棕壤土,试验材料为用贝达、1103P、3309C、140Ru、5C、SO4、华葡1号、抗砧1号8种绿枝嫁接的‘87-1’,砧木于2014年5月定植,2014年6月进行绿枝高接,株行距2.5 m×4.0 m,树形采取单层水平龙干形,叶幕形采取水平叶幕,肥水管理采取水肥一体化,其他管理同常规。
砧木简介：（1）贝达（Beta）,原产于美国,亲本河岸葡萄和美洲葡萄,植株生长势强,抗寒能力强,抗旱性中等,耐盐性中等,耐石灰性土壤中等,我国西北、东北、华北地区主要用作抗寒砧木,扦插生根容易,与大多数品种嫁接亲和性和好。
（2）1103P,原产意大利,亲本为冬葡萄和沙地葡萄,抗根瘤蚜,较抗旱,耐石灰性土壤,耐湿,生长势旺,生根和嫁接状况良好,产枝量中等。
（3）3309C,原产于法国,亲本为河岸葡萄和沙地葡萄,抗根瘤蚜性能优良,抗根癌病也强,根系抗寒能力和抗旱能力中等,耐石灰性土壤中等,生长势中庸,易生根,易嫁接。
（4）140Ru,原产于意大利,亲本为冬葡萄和沙地葡萄,根系抗根瘤蚜,但可能在叶片上面带有虫瘿,抗线虫能力强,耐石灰性土壤,抗旱性中等,不耐湿,较耐酸。
（5）5C,原产于法国,亲本为冬葡萄和河岸葡萄,
抗根瘤蚜,抗线虫病,抗旱,耐寒,耐湿,耐石灰性土壤能力强。长势中等旺盛,根系分布中深,新梢生长快,扦插生根能力中等。
（6）S04,原产于德国,亲本为河岸葡萄和河岸葡萄,抗根瘤蚜,高抗根癌病,抗根结线虫,抗旱性较强,耐湿性强,很耐酸,耐石灰性土壤。
（7）华葡1号（HPYH）,中由国农业科学院果树研究所2011年育成,亲本为白马拉加和左山一,抗寒性强、易生根,树势中庸,植株生长势强。嫁接亲和力好。一年生成熟枝条红褐色,嫩梢绿色。幼叶黄绿色。
（8）抗砧1号（KZYH）,中国农业科学院郑州果树研究所2009年育成,亲本为河岸葡萄和SO4,树势较强,抗根结线虫,嫁接亲和力强。

1.2 试验方法

1.2.1 表观量子效率、光补偿点、暗呼吸速率的测定每个砧穗组合选择长势一致的葡萄3株,每株树挑选其中长势较为一致的带花穗健壮枝条,于晴朗无云的上午,利用Li-6400光合仪,选择其最佳功能叶进行测量（最佳功能叶确定：统一时间段,测定健壮枝条不同节位葡萄叶片瞬时光合速率,选择光合速率最大的叶片作为最佳功能叶,试验得知,最佳功能叶一般位于5—6节位）。设定CO₂浓度为400 μmol·mol^-1,温度为25℃,气体流速为500 mmol·s^-1。设定光合有效辐射（photosynthetically active radiation, PAR）按照由强到弱的顺序分别为2 000、1 800、1 500、1 200、800、400、200、100、50、20、0 μmol·m^-2·s^-1,测定光响应曲线。利用直角双曲线修正模型求得光补偿点（light compensation point,LCP）、暗呼吸速率（dark respiratory rate, Rd）、初始表观量子效率（apparent quantum yield,AQY）^[15]。直角双曲线修正模型表达式如下：

式中,P_n：净光合速率（μmol?m^-2?s^-1）,α：初始量子效率;I：光量子通量密度（μmol?m^-2?s^-1）;R_d：植物的暗呼吸速率（μmol?m^-2?s^-1）,β和γ为系数。
1.2.2 羧化效率、CO₂补偿点的测定设定光合有效辐射（photosynthetically active radiation,PAR）为1 200 μmol·m^-2·s^-1,温度和气体流速分别设为25℃和500 mmol·s^-1。CO₂浓度按照400、200、100、50、20、400、400、800、1 200、1 500、1 800、2 000 μmol·mol^-1的顺序进行测定,利用直角双曲线修正模型拟合CO₂响应曲线,并求得CO₂补偿点（CO₂ compensation point,CCP）和羧化效率（carboxylation efficiency,CE）^[15],模型表达式同上。
1.2.3 高温下净光合速率及不同温度下光合速率变化值测定设定光合有效辐射（photosynthetically active radiation,PAR）为1 200 μmol·m^-2·s^-1,CO₂浓度控制为400 μmol·mol^-1,气体流速为500 mmol·s^-1。温度按照由小到大的顺序设定为25、27、30、32、35和37℃,测定不同温度下净光合速率（net photosynthetic rate,NPR）,并计算不同温度下光合速率变化值。
1.2.4 叶绿素荧光的测定采用英国Hansatech公司的FMS-2型便携脉冲调制式荧光仪测定。每株树均使用最佳功能叶进行测定。在树体直接测定,叶片先暗适应30 min,然后测定暗适应下的荧光参数,其中F₀为初始荧光,Fm为最大荧光,Fv/Fm=(Fm-F₀)/Fm。用弱测量光测量初始荧光（F₀）,然后给一个强闪光（5 000 μmol·m^-2·s^-1）,脉冲时间0.7 s,测得最大荧光（Fm）。
1.2.5 Topsis综合评价法 Topsis法是一种适用于根据多项指标、对多个方案进行比较选择的分析方法,能够客观全面地反映目标状况的动态变化,通过在目标空间中定义一个测度,以此测量目标靠近正理想解和远离负理想解的程度来评估目标的绩效水平,该分析方法可以充分利用原有的数据信息^[16]。本试验中,权重确定由行内专家共同讨论确定,所以只需要根据此权重,确定正负理想解和距离即可。第1步,确定正负理想解：

式中,Y⁺：最偏好的方案（正理想解）,Y^-：最不偏好的方案（负理想解）。
第2步,计算距离。分别计算每个砧穗组合评价向量到正理想解的距离D⁺和负理想解的距离D^-。

式中,贴近度C_j的值介于0—1,C_j越大,表明第j个砧穗组合环境适应性越接近最优水平。

1.3 数据分析方法

采用SAS9.4进行分析,显著性分析采用邓肯新复极差法。

2 结果

2.1 不同砧木对‘87-1’表观量子效率、光补偿点、暗呼吸速率的影响

表观量子效率、光补偿点、暗呼吸速率可以反映不同砧穗组合的耐弱光能力。由表1可以看出,不同砧穗组合表观量子效率、光补偿点、暗呼吸速率均有差异,2016年,嫁接在华葡1号、3309C、SO4、5C、贝达的5种砧木的‘87-1’叶片的表观量子效率显著高于嫁接在140Ru、1103P、抗砧1号3种砧木的。其中‘87-1’/3309C组合光补偿点最低,‘87-1’/5C组合的光补偿点显著高于其他砧穗组合;‘87-1’/5C组合暗呼吸速率最高,而‘87-1’/3309C、‘87-1’/140Ru、‘87-1’/1103P、‘87-1’/抗砧1号、‘87-1’/贝达暗呼吸速率较低。2017年测定的结果有所变化,‘87-1’/SO4组合的表观量子效率显著高于其他组合,‘87-1’/抗砧1号组合的光补偿点显著高于其他组合,而嫁接在SO4、1103P及3309C上的‘87-1’叶片光补偿点较低。‘87-1’/3309C、‘87-1’/抗砧1号暗呼吸速率显著高于其余组合。经过Topsis综合评价法对两年的数据进行计算得分排名,最终得到‘87-1’/3309C组合得分最高,耐弱光能力最强。
Table 1
表1
表1不同砧穗组合表观量子效率、光补偿点、暗呼吸速率
Table 1The apparent quantum yield, light combination point and dark respiration rate of different rootstock-scion combination

砧穗组合 Rootstock-scion combination	年份 Years	表观量子效率 Apparent quantum yield	光补偿点 Light combination point (μmol·m^-2·s^-1)	暗呼吸速率 Dark respiration rate (μmol·m^-2·s^-1)	Topsis得分 Topsis score		Topsis排名 Topsis ranking
砧穗组合 Rootstock-scion combination	年份 Years	表观量子效率 Apparent quantum yield	光补偿点 Light combination point (μmol·m^-2·s^-1)	暗呼吸速率 Dark respiration rate (μmol·m^-2·s^-1)	得分 Score	总分 Total score	Topsis排名 Topsis ranking
‘87-1’/HPYH	2016	0.0758a	23.309bc	1.5222b	0.5862	1.0433	4
‘87-1’/HPYH	2017	0.0657b	23.4195c	0.7787d	0.4571	1.0433	4
‘87-1’/3309C	2016	0.0778a	17.5864d	1.2661c	0.3614	1.2148	1
‘87-1’/3309C	2017	0.0694b	19.3291de	1.9137a	0.8534	1.2148	1
‘87-1’/140Ru	2016	0.0625b	22.4564bc	1.3096c	0.2225	0.7233	7
‘87-1’/140Ru	2017	0.0563c	27.1271b	1.3909b	0.5008	0.7233	7
‘87-1’/1103P	2016	0.0624b	20.3795cd	1.1497c	0.5394	1.194	2
‘87-1’/1103P	2017	0.0439d	18.3896ef	0.7745d	0.6546	1.194	2
‘87-1’/SO4	2016	0.0713a	23.489abc	1.5487b	0.6334	0.0337	5
‘87-1’/SO4	2017	0.0987a	15.7004f	1.367b	0.4003	0.0337	5
‘87-1’/KZYH	2016	0.0573b	25.2786ab	1.3098c	0.2472	0.6468	8
‘87-1’/KZYH	2017	0.0698b	31.186a	1.9143a	0.3996	0.6468	8
‘87-1’/5C	2016	0.0799a	26.8487a	1.9114a	0.4063	0.7399	6
‘87-1’/5C	2017	0.0661b	22.7534dc	1.177c	0.3336	0.7399	6
‘87-1’/Beta	2016	0.0745a	18.9303d	1.3077c	0.3501	1.1017	3
‘87-1’/Beta	2017	0.0548c	22.8865c	1.1474c	0.7516	1.1017	3

Different smalls letters indicated significant difference at 0.05 level by Duncan’s test. The same as below不同小写字母表示差异显著（Duncan,P<0.05）。下同

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2.2 不同砧木对‘87-1’葡萄羧化效率、CO₂补偿点的影响

羧化效率、CO₂补偿点可以反映不同砧穗组合的耐低浓度CO₂能力。由表2可知,2016年,‘87-1’/贝达组合羧化效率显著高于其他组合,‘87-1’/SO4和‘87-1’/5C组合CO₂补偿点显著高于其他组合,‘87-1’/华葡1号和87-1/3309C组合CO₂补偿点最低。2017年,‘87-1’/3309C、‘87-1’/1103P、‘87-1’/140Ru组合表观量子效率显著高于其他组合,‘87-1’/抗砧1号CO₂补偿点显著高于其他组合,‘87-1’/3309C、‘87-1’/华葡1号、‘87-1’/140Ru等组合CO₂补偿点显著低于其他组合。两年数据均表明,‘87-1’/华葡1号和‘87-1’/3309C组合CO₂补偿点较低。另外,无论是表观量子效率、CO₂补偿点,2017年数据都较2016年数据偏高。运用Topsis综合评价,发现‘87-1’/3309C组合得分最高,耐低浓度CO₂能力最强。
Table 2
表2
表2不同砧穗组合羧化效率、CO₂补偿点
Table 2The carboxylation efficiency, CO₂combination point of different rootstock-scion combination

砧穗组合 Rootstock-scion combination	年份 Year	羧化效率 Carboxylation efficiency	CO₂补偿点 CO₂ combination point (μmol?m^-2?s^-1)	Topsis得分 Topsis score		Topsis排名 Topsis ranking
砧穗组合 Rootstock-scion combination	年份 Year	羧化效率 Carboxylation efficiency	CO₂补偿点 CO₂ combination point (μmol?m^-2?s^-1)	得分 Score	总分 Total score	Topsis排名 Topsis ranking
‘87-1’/HPYH	2016	0.0391cd	74.6669d	0.5677	1.0955	2
‘87-1’/HPYH	2017	0.087ab	77.2568c	0.5278	1.0955	2
‘87-1’/3309C	2016	0.0615ab	72.2445d	0.5465	1.1132	1
‘87-1’/3309C	2017	0.0894a	77.2742c	0.5667	1.1132	1
‘87-1’/140Ru	2016	0.0331d	86.5089b	0.5308	0.9881	4
‘87-1’/140Ru	2017	0.1002a	80.0318c	0.4573	0.9881	4
‘87-1’/1103P	2016	0.0554ab	79.4095c	0.5203	1.0279	3
‘87-1’/1103P	2017	0.0913a	85.53c	0.5076	1.0279	3
‘87-1’/SO4	2016	0.0267d	95.7802a	0.4525	0.9059	7
‘87-1’/SO4	2017	0.0408d	96.2671b	0.4534	0.9059	7
‘87-1’/KZYH	2016	0.0532ab	86.0179b	0.4659	0.9238	6
‘87-1’/KZYH	2017	0.0291d	109.7934a	0.4579	0.9238	6
‘87-1’/5C	2016	0.0508bc	96.0808a	0.4331	0.8197	8
‘87-1’/5C	2017	0.0749b	97.0057b	0.3866	0.8197	8
‘87-1’/Beta	2016	0.0649a	79.12c	0.4389	0.9742	5
‘87-1’/Beta	2017	0.0565c	84.6062c	0.5353	0.9742	5

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2.3 不同砧木对‘87-1’高温下净光合速率及不同温度下光合速率变化值的影响

不同砧木对‘87-1’耐高温能力的影响可以通过高温下净光合以及不同温度下净光合速率变化值来反应。由于Li-6400光合仪自身限制,只能测定外界温度±7℃以内的净光合速率值,2016花期进行测定时,外界温度28℃,因此最高温度只能设为35℃,而2017年花期进行测定时,外界温度为30℃,因此温度上限可以设为37℃。有许多研究表明葡萄的最适宜生长温度为25—30℃,超过30℃,光合速率迅速下降^[17,18],所以温度起点设为25℃。由表3可知,对于2016年测定值,‘87-1’/3309C组合35℃净光合速率最大,‘87-1’/5C组合不同温度下净光合速率变化值最小,对于2017年测定值,‘87-1’/1103P组合37℃净光合速率最大,‘87-1’/SO4组合不同温度下净光合速率变化值最小,经Topsis综合排名可知,‘87-1’/SO4得分最高,耐高温能力最强。
Table 3
表3
表3不同砧穗组合不同温度条件下的光合速率
Table 3Net photosynthetic rate in different temperature of different rootstock-scions combination

砧穗组合 Rootstock-scion combination	年份 Year	25℃	27℃	30℃	32℃	35℃	37℃	ΔX	Topsis得分 Topsis score		Topsis排名 Topsis ranking
砧穗组合 Rootstock-scion combination	年份 Year	25℃	27℃	30℃	32℃	35℃	37℃	ΔX	得分 Score	总分 Total score	Topsis排名 Topsis ranking
‘87-1’/HPYH	2016	12.4	12.2	12.2	12.0	11.8	—	0.6	0.4052	1.717	2
‘87-1’/HPYH	2017	17	16.5	16	14.9	13.9	13.1	3.9	0.6066	1.717	2
‘87-1’/3309C	2016	14.50	14.8	14.5	14.3	14.0	—	0.8	0.3204	1.0118	6
‘87-1’/3309C	2017	17	15.7	14.6	12.7	11.9	11.3	5.7	0.3807	1.0118	6
‘87-1’/140Ru	2016	13.9	13.7	13.5	13.0	12.8	—	1.1	0.199	0.8932	7
‘87-1’/140Ru	2017	16	14.2	13	12.4	11.9	11.3	4.7	0.4297	0.8932	7
‘87-1’/1103P	2016	13.0	12.6	12.2	11.7	11.4	—	1.6	0.1033	0.892	5
‘87-1’/1103P	2017	18.9	18.3	17.4	16.3	15.8	14.6	4.3	0.6105	0.892	5
‘87-1’/SO4	2016	14.3	14.2	13.9	13.6	13.3	—	1	0.2353	0.7138	1
‘87-1’/SO4	2017	16.7	16.2	15.4	14.8	14.5	14	2.7	0.901	0.7138	1
‘87-1’/KZYH	2016	13.8	13.4	13.20	12.7	12.3	—	1.5	0.143	0.7011	8
‘87-1’/KZYH	2017	9.5	9.32	8.25	7.25	6.02	5.61	3.89	0.2439	0.7011	8
‘87-1’/5C	2016	10.99	11.09	10.90	10.87	10.64	—	0.56	0.4559	0.6287	4
‘87-1’/5C	2017	10.2	8.42	7.83	7.76	7.24	7.11	3.09	0.4361	0.6287	4
‘87-1’/Beta	2016	12.5	12.30	12.0	11.60	11.40	—	1.1	0.157	0.3869	3
‘87-1’/Beta	2017	16.2	15.1	13.8	13.8	13.3	13	3.2	0.7362	0.3869	3

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2.4 不同砧木对‘87-1’叶绿素荧光特性的影响

叶绿素荧光特征能够反映植株叶片的光合效率和潜在能力^[19]。由表4可知,‘87-1’/140Ru和‘87-1’/SO4组合初始荧光F₀最高,但与‘87-1’/1103P、‘87-1’/5C、‘87-1’/抗砧1号3个组合无显著性差异。而‘87-1’/1103P组合可变荧光Fv最高,显著高于‘87-1’/抗砧1号、‘87-1’/贝达和‘87-1’ /华葡1号3个组合,与剩余组合无显著性差异。对于最大荧光Fm和PSII最大光化学效率Fv/Fm而言,所有砧穗组合均无显著性差异。此外,‘87-1’/1103P和‘87-1’/3309C 2个组合的Fv/F₀显著高于其他组合。但整体而言,不同砧穗组合的叶绿素荧光参数相差不大。
Table 4
表4
表4不同砧穗组合的叶绿素荧光参数
Table 4Chlorophyll fluorescence parameters of different rootstock-scions combinations

砧穗组合 Rootstock-scion combination	F₀	Fv	Fm	Fv/Fm	Fv/F₀
‘87-1’/HPYH	233.0b	1003bc	1236a	0.811a	4.301ab
‘87-1’/3309C	232.7b	1055.7abc	1288a	0.817a	4.529a
‘87-1’/140Ru	257.7a	1099ab	1295a	0.771a	4.264ab
‘87-1’/1103P	248ab	1149.7a	1217a	0.811a	4.689a
‘87-1’/SO4	253.3a	1095.7ab	1349a	0.812a	4.345ab
‘87-1’/KZYH	237.7ab	944c	1181a	0.798a	3.970b
‘87-1’/5C	244.7ab	1021.3abc	1266a	0.807a	4.176ab
‘87-1’/Beta	230.7b	969.7bc	1200a	0.807a	4.201ab

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2.5 不同砧穗组合光合参数和叶绿素荧光参数的相关性分析

不同砧穗组合的Fv/F₀值与光补偿点呈显著性负相关,与高温下净光合速率成显著性正相关,其光补偿点与高温下净光合速率呈显著性负相关,羧化效率和CO₂补偿点成显著性正相关。
Table 5
表5
表5不同砧穗组合光合参数和叶绿素荧光参数的相关系数
Table 5The correlation coefficient of photosynthetic parameters and chlorophyll fluorescence parameters of different rootstock- scion combinations

	F₀	Fv	Fm	Fv/Fm	Fv / F₀	表观量子效率 AQY	光补偿点 LCP	暗呼吸速率 Rd	羧化效率 CE	CO₂ 补偿点 CCP	高温下净光合速率 NPR	ΔX
F₀	1
Fv	0.70637	1
Fm	0.57505	0.55952	1
Fv/Fm	-0.55284	-0.06732	-0.046	1
Fv / F₀	0.14838	0.80414	0.27875	0.38603	1
表观量子效率AQY	0.14665	-0.08636	0.63229	0.25166	-0.24755	1
光补偿点LCP	-0.13968	-0.61891	-0.55659	-0.63451	-0.75195*	-0.28537	1
暗呼吸Rd	-0.10423	-0.3034	0.12864	-0.12677	-0.34982	0.38938	0.32475	1
羧化效率CE	0.14869	0.53979	0.17599	-0.18706	0.60889	-0.56644	-0.17226	-0.38793	1
CO₂补偿点CCP	0.14281	-0.36561	-0.20268	-0.01505	-0.60409	0.37939	0.32798	0.36329	-0.82159*	1
高温下净光合速率 NPR	0.12225	0.59286	0.29319	0.25136	0.72352*	-0.07224	-0.71057*	-0.57168	0.3565	-0.65507	1
ΔX	-0.1395	0.24452	-0.01394	-0.12738	0.4371	-0.38358	0.1165	0.36265	0.61024	-0.53241	0.01598	1

*represent significant correlation at the 0.05 level*表示在0.05水平上显著相关

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3 讨论

表观量子效率是指光合作用机构每吸收1 mol光量子后光合释放的O₂摩尔数或同化CO₂的摩尔数^[20,21],LEE等^[22]和BJ?RKMAN等^[23]的研究认为耐荫植物具有较高表观量子效率。表观量子效率越高,说明叶片光能转化效率越高,对弱光的利用率越强。光补偿点反映了植物叶片光合作用过程中光合同化作用与呼吸消耗相当时的光强。光补偿点低说明植物利用弱光能力强,有利于有机物质的积累^[24]。暗呼吸速率反映了在弱光下,消耗有机物的多少,一般而言,暗呼吸速率越低,耐弱光能力越强。以上3个指标可以反映不同砧穗组合的耐弱光能力。由本试验可知,‘87-1’/3309C和‘87-1’/1103P组合耐弱光能力较强。CO₂补偿点是维持植物生长的最低CO₂浓度,代表植株开始累积同化物起始点^[25],而羧化效率反映低CO₂浓度下植物Rubisco羧化氧化酶的活性大小^[26],以上两个指标可以反映不同砧穗组合耐低浓度CO₂的能力。由本试验可知,‘87-1’/3309C和‘87-1’/华葡1号耐低浓度CO₂能力较强。高温下净光合速率是指在光合仪可以设定的最高温下测定的净光合速率的值,不同温度下净光合速率变化值指从25℃到最高温之间净光合速率的差值。以上两者可以反映不同砧穗组合耐高温能力。本试验中‘87-1’/SO4、‘87-1’/华葡1号耐高温能力较强。
叶绿素荧光可以作为光合作用的有效探针,能够反映一些光合生理的重要指标^[27,28,29],Fv/Fm是指经过充分暗适应的植物叶片PSII最大光化学效率,是表示PSII光化学效率的重要指标。其大小反映了PSII反应中心内原初光能的转化效率,反映植物潜在的最大光合能力。Fv/Fm值越大,表明该植物的光能利用潜力越大^[30,31,32]。而本试验中8种砧穗组合的Fv/Fm并无显著性差异,说明这8种砧木对‘87-1’葡萄的最大光合能力影响不大。Fv/F₀代表PSII的潜在光化学活性,反映原初光能转化效率及PSII潜在量子效率。本试验发现87-1/1103P和87-1/3309C 2个组合Fv/F₀最高,说明1103P和3309C做砧木有效保护了葡萄叶片放氧复合体和反应中心,使PSII维持较高的光化学活性^[18]。此外,经相关分析可知,F_V/F₀与光补偿点呈显著性负相关,与高温下净光合速率成显著性正相关,说明Fv/F₀高的砧穗组合耐弱光能力和耐高温能力强。‘87-1’/140Ru和‘87-1’/SO4组合初始荧光F₀最大,‘87-1’/1103P组合可变荧光Fv最高,F₀的增加可能是植物叶片PSII反应中心出现可逆的失活或出现不易逆转的破坏,也可能是植物叶片类囊体膜受到损伤,而且F₀增加量越多,类囊体膜受损程度就越严重。类囊体膜结构发生改变,首先反映的是初始荧光F₀的上升。对于具有高初始荧光F₀的‘87-1’/140Ru和‘87-1’/SO4组合,说明两者对外界环境变化敏感,叶片类囊体膜易受到损伤。
不同砧木会对‘87-1’的光合特性和叶绿素荧光特性会产生影响,主要是由砧木遗传特性决定的。3309C由河岸葡萄和沙地葡萄杂交而成,生根性能和嫁接亲和性都很好,耐湿、抗病;140Ru、1103P由冬葡萄和沙地葡萄杂交而来,使接穗新梢生长势强、产量高。国内外研究也表明,1103P较其他砧木有较稳定的光合能力^[33,34]。5C、SO4由河岸葡萄和冬葡萄杂交而来,易生根,嫁接亲和性好^[35]。华葡1号由白马拉加和左山一杂交而来,抗寒、易生根,树势中庸,但不抗葡萄根瘤蚜。抗砧1号由河岸葡萄和SO4杂交而来,树势较强。贝达由美洲葡萄和河岸葡萄杂交而来,嫁接亲和性好,抗寒、抗涝能力强,对根瘤蚜敏感。所以,可以初步推测,易生根嫁接亲和性好的砧木有利于提高设施栽培‘87-1’耐弱光能力、耐低浓度CO₂能力、耐高温能力。新梢生长势强的冬葡萄和沙地葡萄杂交后代PSⅡ的潜在活性和原初光能转化效率高。至于其中的机理,还需要进一步研究。此外,不同砧木产生的植物激素、生理活性物质和特殊代谢产物种类和数量有所差异,并且对营养物质运输、吸收、同化和利用能力不同,从而导致了接穗光合性能的差异。所以,在选择砧木时,要根据当地的气候环境,综合考虑各种因素,选择适宜当地自然环境的砧木进行嫁接种植,最大限度的发挥利用砧木的自身遗传特性,改善接穗的生长状况。对于设施葡萄而言,最关注的就是耐弱光、耐低CO₂浓度、耐高温的能力^[36],当然,抗根瘤蚜能力是砧木筛选的前提。

4 结论

3309C、1103P砧木可以有效提高‘87-1’耐弱光能力及原初光能转化效率,3309C、华葡1号砧木可以有效提高‘87-1’的耐低浓度CO₂能力,SO4、华葡1号砧木可以有效提高‘87-1’的耐高温能力。3309C、SO4、1103P 3种砧木可以考虑作为设施专用砧木备选品种,生产中可以根据实际需求相应选择。
The authors have declared that no competing interests exist.

参考文献原文顺序
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被引期刊影响因子

[1]	VRSIC S, PULKO B, KOCSIS L.Factors influencing grafting success and compatibility of grape rootstocks .Scientia Horticulturae, 2015, 181: 168-173.https://doi.org/10.1016/j.scienta.2014.10.058 URL [本文引用: 2]摘要 The aim of this work was to examine the compatibility of different rootstocks with various wine grape varieties. In the first trial, Vitis vinifera L. cvs. Furmint, Regent, Riesling, Sauvignon Blanc, and Welschriesling were grafted onto B02rner, 5BB and SO4 rootstocks. In the second trial Welschriesling was grafted on 12 rootstocks. Their growth during the first year in the vineyard was measured. As an index of compatibility, the percentage of first grade grafted vines as well as dry weight of mature shoots (canes) and roots was determined. After the grafts went through the first phase of healing often called callusing (2–3 weeks of moist warm conditions), the differences in callus formation were greater between years than among rootstocks, which were the most obvious with Furmint as a scion. No good callus formation occurred when cane (rootstock, scion) maturity is not adequate. Such was the case in 2005, there were 24% fewer grafts with fully developed callus than those in 2006. The dry weight of roots was higher on 5BB than on B02rner and SO4. B02rner rootstock had fewer roots, and the roots were thinner. Welschriesling had good compatibility with all rootstocks (the average grafts success was 67%), but that of 5BB, G251, and G103 (above 80%) was greater than the average. The G103 rootstock had the highest dry weight of roots. Shoot growth in vineyards was above the average with 5BB, SO4, Binova, B02rner, and M V rootstocks. All Georgikon rootstocks had a lower cane dry weight per vine than the others.
[2]	JIN Z X, SUN H, SUN T Y, WANG Q J, YAO Y X.Modifications of ‘Gold Finger’ grape berry quality as affected by the different rootstocks .Journal of Agricultural and Food Chemistry, 2016, 64: 4189-4197.https://doi.org/10.1016/j.scienta.2016.07.023 URLPMID:27088562 [本文引用: 1]摘要 Grafting grapevines on resistant rootstocks is an effective tool to overcome biotic and abiotic stress constraints. The increasing consumer attention for fruit quality makes it indispensable to evaluate the impacts of resistant rootstocks on fruit quality. In this paper, berry quality modifications induced by the rootstocks were determined in the ‘Summer Black’ grape. Compared to the own-rooted vines, berry and cluster weights, berry texture and skin color were altered by the rootstocks to varying extents. Beta maintained TSS/TA and the contents of fructose, glucose and sucrose; in contrast, SO4, 5BB and 101–14M impaired these parameters. The rootstocks modified berry nutrition and flavor by affecting the contents of cations and free amino acids. Aroma amount and components were largely modified by the rootstocks; Beta largely enhanced the amount of total aroma volatiles and ester compounds while SO4 imparted the reverse influence. Hence, berry nutrition and flavor were widely modified by the rootstocks; the Beta rootstock favored berry quality, and SO4 impaired berry quality; the effects of 5BB and 101–14M on berry quality varied according to the different parameters.
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[7]	杨瑞. 葡萄砧穗组合筛选及嫁接早期亲和力研究[D] . 兰州: 甘肃农业大学, 2007. [本文引用: 1] YANG R.Selections stock-scion combination and studies on graft affinity in early stage of different combination of rootstocks and scions of grape [D] . Lanzhou: Gansu Agricultural University, 2007 (in Chinese) [本文引用: 1]
[8]	SALIMIA R B, HAMDAN S.GRAPEVINE scion-rootstock combinations of palestinian local cultivars and rootstocks resistant to grape phylloxera daktulosphaira vitifoliae (Fitch) [Phylloxeridae: Homoptera] .Dirasat Agricultural Sciences, 2009(1): 19-28. [本文引用: 1]
[9]	LI Z, MARGUERIT E.The influence of grapevine rootstocks on scion growth and drought resistance .Theoretical and Experimental Plant Physiology, 2016, 28(2): 1-15.https://doi.org/10.1007/s40626-016-0070-x URL [本文引用: 1]摘要 Grapes are a widely cultivated and economically important crop. Climate change is increasing the focus and investment on the development of more drought resistant varieties. However, markets often dic
[10]	李敏敏, 袁军伟, 刘长江,韩斌, 黄家珍, 郭紫娟, 赵胜建. 砧木对河北昌黎产区赤霞珠葡萄生长和果实品质的影响 . 应用生态学报, 2016, 27(1): 59-63.https://doi.org/10.13287/j.1001-9332.201601.003 URL Magsci [本文引用: 1]摘要以7个砧木嫁接的赤霞珠和自根为材料,研究不同砧木对河北昌黎产区赤霞珠葡萄生长、结果和品质的影响.结果表明:5C和5BB显著提高赤霞珠主干粗度,5C显著提高赤霞珠主梢粗度;188-08、5BB和5C显著提高赤霞珠可溶性固形物含量,188-08和5BB显著提高还原糖含量;101-14M和3309C显著降低可滴定酸含量,5C和101-14M显著提高果皮多酚和花青素含量,101-14M显著提高果皮单宁含量;3309C、110R、5C和101-14M显著提高赤霞珠单株产量.采用模糊评价法综合评价7个砧木对赤霞珠生长、品质和产量等因子的影响,在河北昌黎地区独特的气候和环境条件下,7种砧木嫁接赤霞珠的综合评价均优于自根,适宜砧木顺序依次为5C、101-14M、3309C、5BB、188-08、110R、SO4. LI M M, YUAN J W, LIU C J, HAN B, HUANG J Z, GUO Z J, ZHAO S J.Effects of rootstocks on the growth and berry quality of Vitis vinifera cv. Cabernet Sauvignon grapevine in Changli zone, Hebei province, China. Chinese Journal of Applied Ecology, 2016, 27(1): 59-63. (in Chinese)https://doi.org/10.13287/j.1001-9332.201601.003 URL Magsci [本文引用: 1]摘要以7个砧木嫁接的赤霞珠和自根为材料,研究不同砧木对河北昌黎产区赤霞珠葡萄生长、结果和品质的影响.结果表明:5C和5BB显著提高赤霞珠主干粗度,5C显著提高赤霞珠主梢粗度;188-08、5BB和5C显著提高赤霞珠可溶性固形物含量,188-08和5BB显著提高还原糖含量;101-14M和3309C显著降低可滴定酸含量,5C和101-14M显著提高果皮多酚和花青素含量,101-14M显著提高果皮单宁含量;3309C、110R、5C和101-14M显著提高赤霞珠单株产量.采用模糊评价法综合评价7个砧木对赤霞珠生长、品质和产量等因子的影响,在河北昌黎地区独特的气候和环境条件下,7种砧木嫁接赤霞珠的综合评价均优于自根,适宜砧木顺序依次为5C、101-14M、3309C、5BB、188-08、110R、SO4.
[11]	SOUMKUWAR R G, TAWARE P B, BHANGE M A, SHARMA J, KHAN I.Influence of different rootstocks on growth, photosynthesis, biochemical composition, and nutrient contents in‘Fantasy Seedless’ grapes. International Journal of Fruit Science, 2015, 15(3): 1-16.https://doi.org/10.1080/15538362.2015.1031564 URL [本文引用: 1]摘要 The influence of 10 different rootstocks was investigated for evaluating the most suitable rootstock for table grape cultivar Fantasy Seedless. The growth performance, photosynthesis, biochemical composition, and primary nutrients090005 status of the scion after grafting were investigated in 090004Fantasy Seedless090005 grafted on 10 different rootstocks. The studies on vegetative parameters revealed that the rootstock influences the vegetative growth thereby increasing the photosynthesis of a vine. The rootstock influenced the changes in biochemical constituents in the grafted vine thus helping the vine to store enough food material. The grafted vines exhibited variations in primary nutrients status highlighting the scope for selection of better rootstock for sustainable nutrition management.
[12]	李小红, 周凯, 谢周, 李辉信, 章镇, 陶建敏. 不同葡萄砧木对矢富罗莎葡萄嫁接苗光合作用的影响 . 果树学报, 2009, 26(1): 90-93.URL [本文引用: 1]摘要在避雨条件下,以盆栽试验观察不同砧木(巨峰、1202、5BB、110R、99R、华佳8号、8B和SO4)对矢富罗莎葡萄叶片光合日变化的影响。结果表明,不同葡萄砧木矢富罗莎嫁接苗叶片净光合速率(Pn)的日变化基本呈双峰型曲线。叶片净光合速率日变化存在差异,99R、华佳8号和1202砧穗组合嫁接苗叶片的净光合速率(Pn)明显高于其他组合,最高峰值(11:00时)分别高出对照128.8%、106.0%和77.3%;叶片气孔导度(Gs)值1202、110R、99R和华佳8号组合较高,9:00时分别达到对照的8.5、8.0、7.5和6.5倍;99R、华佳8号、110R和1202组合的叶片蒸腾速率(Tr)显著高于其他组合和对照,巨峰和8B组合居中。 LI X H, ZHOU K, XIE Z, LI H X. ZHANG Z, TAO J M.Photosynthesis of Yatomi Rosa on eight grape rootstocks .Journal of Fruit Science, 2009, 26(1): 90-93. (in Chinese)URL [本文引用: 1]摘要在避雨条件下,以盆栽试验观察不同砧木(巨峰、1202、5BB、110R、99R、华佳8号、8B和SO4)对矢富罗莎葡萄叶片光合日变化的影响。结果表明,不同葡萄砧木矢富罗莎嫁接苗叶片净光合速率(Pn)的日变化基本呈双峰型曲线。叶片净光合速率日变化存在差异,99R、华佳8号和1202砧穗组合嫁接苗叶片的净光合速率(Pn)明显高于其他组合,最高峰值(11:00时)分别高出对照128.8%、106.0%和77.3%;叶片气孔导度(Gs)值1202、110R、99R和华佳8号组合较高,9:00时分别达到对照的8.5、8.0、7.5和6.5倍;99R、华佳8号、110R和1202组合的叶片蒸腾速率(Tr)显著高于其他组合和对照,巨峰和8B组合居中。
[13]	刘廷松, 李桂芬. 设施栽培条件下葡萄盛花期的光合特性 . 园艺学报, 2003, 30(5): 568-570.URL Magsci [本文引用: 1]摘要 <FONT face=Verdana>对设施栽培条件下‘红双味’葡萄花期净光合速率(Pn) 日变化规律, 设施内各生态因子,尤其是光照强度和CO<SUB>2</SUB> 浓度对葡萄光合作用的影响进行了研究。晴天栽培设施内葡萄叶片的Pn 日变化呈明显的双峰曲线, 阴天时葡萄叶片Pn 的日变化趋势主要决定于光照强度的日变化。晴天的Pn 明显高于阴天。研究葡萄叶片Pn 对光照强度和CO<SUB>2 </SUB>浓度的响应曲线得出, 葡萄叶片的光饱和点为1000μmol·m<SUP>-2</SUP>·s <SUP>-1</SUP> , 光补偿点为21.3μmol·m<SUP>-2</SUP>·s <SUP>-1</SUP> , CO<SUB>2</SUB> 饱和点为700μL·L <SUP>-1</SUP> , CO2 补偿点为54.6μL·L<SUP> -1</SUP> 。</FONT> LIU Y S, LI G F.Some photosynthetic characteristics of blooming grape in protected cultivation .Acta Horticulturae Sinica, 2003, 30(5): 568-570. (in Chinese)URL Magsci [本文引用: 1]摘要 <FONT face=Verdana>对设施栽培条件下‘红双味’葡萄花期净光合速率(Pn) 日变化规律, 设施内各生态因子,尤其是光照强度和CO<SUB>2</SUB> 浓度对葡萄光合作用的影响进行了研究。晴天栽培设施内葡萄叶片的Pn 日变化呈明显的双峰曲线, 阴天时葡萄叶片Pn 的日变化趋势主要决定于光照强度的日变化。晴天的Pn 明显高于阴天。研究葡萄叶片Pn 对光照强度和CO<SUB>2 </SUB>浓度的响应曲线得出, 葡萄叶片的光饱和点为1000μmol·m<SUP>-2</SUP>·s <SUP>-1</SUP> , 光补偿点为21.3μmol·m<SUP>-2</SUP>·s <SUP>-1</SUP> , CO<SUB>2</SUB> 饱和点为700μL·L <SUP>-1</SUP> , CO2 补偿点为54.6μL·L<SUP> -1</SUP> 。</FONT>
[14]	綦伟, 厉恩茂, 翟衡, 王晓芳, 杜远鹏, 谭皓. 部分根区干旱对不同砧木嫁接玛瓦斯亚葡萄生长的影响 . 中国农业科学, 2007, 40(4): 794-799.https://doi.org/10.3321/j.issn:0578-1752.2007.04.020 URL Magsci [本文引用: 1]摘要 Biological response of the sane grape cultivar grafted by different rootstocs to the different irrigation metods were studied, which provided biological basement for spreading and using resistant rootstocks and saving-water irrigation. Biological effects of 1/2 divided root irrigation on M/420A、M/3309C and M/110Rwere studied by simulating soil boxes with controlled alternate irrigation for half part of rootzone, controlled fixed irrigation for half part of rootzone,and conventional irrigation for whole root zone, and the grafted graoe had been anatomized after the leaf falled. The results showed controlled alternate irrigation remarkably promoted the growth of current-grew roots , new roots numbers, the surface area and volume of effective roots and roots length of the three kinds cultivar/rootstock, however restricted the growth of new shoots compared with conventional irrigation. Then root-shoot ratio of M/3309C、M/420A and M/110R increased 41.18%、23.68% and37.5%. Under controlled fixed irrigation, the growth and length of new roots in dry zone remarkably decreased, M/3309C which drought-resistent was lower reduced especially most compared with CK, and reduced 67.67%,then induced total mass of new roots and growth of new shoots decreased sharply. The prolonged growth of shoot internodes were restricted under controlled alternate irrigation and controlled fixed irrigation, M/110R reduced a little which the base shoot internodes reduced 7.37% and 23.74% respectively and the top shoot internodes reduced 9.72% and 19.75% respectively. The research indicated different cultivar/rootstocks had obvious distinction to adapt to water stress, alternation irrigation contributed to increase roots growth of grape, improve the ability of drought-resistent. QIN W, LI E M, ZHAN H, WANG X F, DU Y P, TAN H.Effects of partial rootzone drying on the growth of Vitis vinifera cv. malvasia grafted on varied rootstocks. Scientia Agricultura Sinica, 2007, 40(4): 794-799. (in Chinese)https://doi.org/10.3321/j.issn:0578-1752.2007.04.020 URL Magsci [本文引用: 1]摘要 Biological response of the sane grape cultivar grafted by different rootstocs to the different irrigation metods were studied, which provided biological basement for spreading and using resistant rootstocks and saving-water irrigation. Biological effects of 1/2 divided root irrigation on M/420A、M/3309C and M/110Rwere studied by simulating soil boxes with controlled alternate irrigation for half part of rootzone, controlled fixed irrigation for half part of rootzone,and conventional irrigation for whole root zone, and the grafted graoe had been anatomized after the leaf falled. The results showed controlled alternate irrigation remarkably promoted the growth of current-grew roots , new roots numbers, the surface area and volume of effective roots and roots length of the three kinds cultivar/rootstock, however restricted the growth of new shoots compared with conventional irrigation. Then root-shoot ratio of M/3309C、M/420A and M/110R increased 41.18%、23.68% and37.5%. Under controlled fixed irrigation, the growth and length of new roots in dry zone remarkably decreased, M/3309C which drought-resistent was lower reduced especially most compared with CK, and reduced 67.67%,then induced total mass of new roots and growth of new shoots decreased sharply. The prolonged growth of shoot internodes were restricted under controlled alternate irrigation and controlled fixed irrigation, M/110R reduced a little which the base shoot internodes reduced 7.37% and 23.74% respectively and the top shoot internodes reduced 9.72% and 19.75% respectively. The research indicated different cultivar/rootstocks had obvious distinction to adapt to water stress, alternation irrigation contributed to increase roots growth of grape, improve the ability of drought-resistent.
[15]	叶子飘. 光合作用对光和CO2响应模型的研究进展 . 植物生态学报, 2010, 34(6): 727-740.https://doi.org/10.3773/j.issn.1005-264x.2010.06.012 URL Magsci [本文引用: 2]摘要 <FONT face=Verdana>光合作用对光和CO<SUB>2</SUB>响应模型是研究植物生理和植物生态学的重要工具, 可为植物光合特性对主要环境因子的响应提供科学依据。该文综述了当前光合作用对光和CO<SUB>2</SUB>响应模型的研究进展和存在的问题, 并在此基础上探讨了这些模型的可能发展趋势。光合作用涉及光能的吸收、能量转换、电子传递、ATP合成、CO<SUB>2</SUB>固定等一系列复杂的物理和化学反应过程。光合作用由原初反应、同化力形成和碳同化3个基本过程构成, 任一个过程均可对光合作用速率产生直接的影响。光合作用对光响应模型只涉及光能的转换, 而光合作用的生化模型包含了同化力形成和碳同化这两个基本过程。把光合作用的原初反应, 即把参与光能吸收、传递和转换的捕光色素分子的物理参数(如捕光色素分子数、捕光色素分子光能吸收截面、捕光色素分子处于激发态的平均寿命等)结合到生化模型中, 可能是今后光合作用对光响应机理模型的发展方向。</FONT> YE Z P.A review on modeling of responses of photosynthesis to light and CO₂ .Chinese Journal of Plant Ecology, 2010, 34(6): 727-740. (in Chinese)https://doi.org/10.3773/j.issn.1005-264x.2010.06.012 URL Magsci [本文引用: 2]摘要 <FONT face=Verdana>光合作用对光和CO<SUB>2</SUB>响应模型是研究植物生理和植物生态学的重要工具, 可为植物光合特性对主要环境因子的响应提供科学依据。该文综述了当前光合作用对光和CO<SUB>2</SUB>响应模型的研究进展和存在的问题, 并在此基础上探讨了这些模型的可能发展趋势。光合作用涉及光能的吸收、能量转换、电子传递、ATP合成、CO<SUB>2</SUB>固定等一系列复杂的物理和化学反应过程。光合作用由原初反应、同化力形成和碳同化3个基本过程构成, 任一个过程均可对光合作用速率产生直接的影响。光合作用对光响应模型只涉及光能的转换, 而光合作用的生化模型包含了同化力形成和碳同化这两个基本过程。把光合作用的原初反应, 即把参与光能吸收、传递和转换的捕光色素分子的物理参数(如捕光色素分子数、捕光色素分子光能吸收截面、捕光色素分子处于激发态的平均寿命等)结合到生化模型中, 可能是今后光合作用对光响应机理模型的发展方向。</FONT>
[16]	WANG TIENCHIN, CHANG TSUNGHAN.Application of TOPSIS in evaluating initial training aircraft under a fuzzy environment .Expert Systems with Applications, 2007, 33(4): 870-880.https://doi.org/10.1016/j.eswa.2006.07.003 URL Magsci [本文引用: 1]摘要 This paper develops an evaluation approach based on the Technique for Order Performance by Similarity to Ideal Solution (TOPSIS), to help the Air Force Academy in Taiwan choose optimal initial training aircraft in a fuzzy environment where the vagueness and subjectivity are handled with linguistic terms parameterised by triangular fuzzy numbers. This study applies the fuzzy multi-criteria decision-making (MCDM) method to determine the importance weights of evaluation criteria and to synthesize the ratings of candidate aircraft. Aggregated the evaluators鈥 attitude toward preference; then TOPSIS is employed to obtain a crisp overall performance value for each alternative to make a final decision. This approach is demonstrated with a real case study involving 16 evaluation criteria, seven initial propeller-driven training aircraft assessed by 15 evaluators from the Taiwan Air Force Academy.
[17]	贺普超. 葡萄学. 北京:中国农业出版社, 1999: 74-75. [本文引用: 1] HE P C. Vitis Culture.Beijing: China Agricultural Press, 1999: 74-75. (in Chinese) [本文引用: 1]
[18]	郑秋玲, 谭伟, 马宁, 翟衡. 钙对高温下巨峰葡萄叶片光合作用和叶绿素荧光的影响 . 中国农业科学, 2010, 43(9): 1963-1968.https://doi.org/10.3864/j.issn.0578-1752.2010.09.025 URL Magsci [本文引用: 2]摘要【Objective】 In order to reveal the protection function of calcium on photosynthetic organ in leaves of ‘Kyoho’ grape under high temperature stress, the effects of CaCl2 on photosynthesis and chlorophyll fluorescence under heat stress were studied. 【Method】 During the period of high temperature, the grapes grown in the field were pretreated with CaCl2, CaCl2+Ca2+ chelator EGTA and CaCl2+CaM antagonist CPZ. Photosynthetic rate, chlorophyll fluorescence and MDA content were determined. 【Result】The net photosynthesis rate (Pn), qP, Fv’/Fm’ and Fv/Fm in the grape leaves pretreated with CaCl2 were much higher than that of the control, while Fo was much less than that of the control, and the accumulation of final product of membrane lipid peroxidation (MDA) was slightly lower than the control. EGTA and CPZ pretreatments significantly inhibited the photosythesis, Pn were 29.4% and 26.5% lower than that of CaCl2 pretreatment; the activity of PSⅡ was inhibited significantly, the accumulation of MDA were 33.7% and 40.2% higher than that of CaCl2 pretreatment. 【Conclusion】 CaCl2 pretreatment effectively alleviated the inhibition of photosynthesis in grape leaves under high temperature stress and the damage to the function of PSⅡ to some extent. ZHENG Q L, TAN W, MA N, ZHAI H.Effects of calcium on photosynthesis and chlorophyll fluorescence of 'Kyoho' grape under high temperature stress .Scientia Agricultura Sinica, 2010, 43(9): 1963-1968. (in Chinese)https://doi.org/10.3864/j.issn.0578-1752.2010.09.025 URL Magsci [本文引用: 2]摘要【Objective】 In order to reveal the protection function of calcium on photosynthetic organ in leaves of ‘Kyoho’ grape under high temperature stress, the effects of CaCl2 on photosynthesis and chlorophyll fluorescence under heat stress were studied. 【Method】 During the period of high temperature, the grapes grown in the field were pretreated with CaCl2, CaCl2+Ca2+ chelator EGTA and CaCl2+CaM antagonist CPZ. Photosynthetic rate, chlorophyll fluorescence and MDA content were determined. 【Result】The net photosynthesis rate (Pn), qP, Fv’/Fm’ and Fv/Fm in the grape leaves pretreated with CaCl2 were much higher than that of the control, while Fo was much less than that of the control, and the accumulation of final product of membrane lipid peroxidation (MDA) was slightly lower than the control. EGTA and CPZ pretreatments significantly inhibited the photosythesis, Pn were 29.4% and 26.5% lower than that of CaCl2 pretreatment; the activity of PSⅡ was inhibited significantly, the accumulation of MDA were 33.7% and 40.2% higher than that of CaCl2 pretreatment. 【Conclusion】 CaCl2 pretreatment effectively alleviated the inhibition of photosynthesis in grape leaves under high temperature stress and the damage to the function of PSⅡ to some extent.
[19]	陈建明, 俞晓平, 程家安. 叶绿素荧光动力学及其在植物抗逆生理研究中的应用 . 浙江农业学报, 2006, 18(1): 51-55.URL [本文引用: 1] CHEN J M, YU X P, CHENG J A.The application of chlorophyll fluorescence kinetics in the study of physiological responses of plants to environmental stresses .Journal of Zhejiang Agricultural University, 2006, 18(1): 51-55. (in Chinese)URL [本文引用: 1]
[20]	HU C, ULLER-KARGER F E, ZEPP R G. MAbsorbance, absorption coefficient, and apparent quantum yield: A comment on common ambiguity in the use of these optical concepts .Limnology & Oceanography, 2002, 47(4): 1261-1267.https://doi.org/10.4319/lo.2002.47.4.1261 URL [本文引用: 1]摘要 Several important optical terms, such as "absorbance" and "absorption coefficient," are frequently used ambiguously in the current peer-reviewed literature. Because these terms are important when deriving other quantities, such as the apparent quantum yield of photoproduction, ambiguity in the application of these concepts leads to results that are difficult or impossible to interpret correctly. Such ambiguity also hinders comparison of results between studies and ultimately harms proper parameterization of numerical models of oceanic processes, as well as the refinement of remote sensing algorithms. We review these concepts and the implications of such ambiguities. A few simple recommendations that follow conventions developed by optical oceanographers are provided to authors dealing with these concepts. In particular, the symbol a is recommended for the absorption coefficient (in Napierian form, m-1), which is also preferred over absorbance (dimensionless) when data are presented; the symbol a is not recommended for absorbance; A should be used with caution because, although it has been used widely for absorbance in photochemistry and photobiology, it has also been used for absorptance in physics and optical oceanography; the term "absorptivity" is not recommended because of conflicting definitions in the current literature; the pathlength should always be given whenever absorbance data are presented; and normalization of photoproduction rates to absorbance or absorption coefficient should be performed only on optically thin samples, unless the inner filter effects are accounted for and corrected.
[21]	XU D Q.Non-uniform stomatal closure and non-stomatal limitation of photosynthesis .Plant Physiology Communications, 1995, 31(4) : 246-252. [本文引用: 1]
[22]	LEE D W, BONE R A, TARSIS S, STORCH D.Correlates of leaf optical properties in tropical forest sun and extreme-shade plants .America Journal of Botany, 1990, 77(3): 370-380.https://doi.org/10.1002/j.1537-2197.1990.tb13566.x URL [本文引用: 1]摘要 Thirteen shade-adapted rain forest species were compared with twelve sun-adapted tropical forest species for correlates to leaf optical properties (described previously in Amer. J. Bot. 73: 1100-1108). The two samples were similar in absorptance of quanta for photosynthesis, but the shade-adapted taxa: 1) had significantly lower specific leaf weights, indicating a more metabolically efficient production of surface for quantum capture; 2) synthesized less chlorophyll per unit area; and 3) used less chlorophyll for capturing the same quanta for photosynthesis. The anatomical features that best correlate with this increased efficiency are palisade cell shape and chloroplast distribution. Palisade cells with more equal dimensions have more chloroplasts on their abaxial surfaces. This dense layer of chloroplasts maximizes the light capture efficiency limited by sieve effects. The more columnar palisade cells of sun-adapted taxa allow light to pass through the central vacuoles and spaces between cells, making chloroplasts less efficient in energy capture, but allowing light to reach chloroplasts in the spongy mesophyll. Pioneer species may be an exception to these two groups of species. Three pioneer taxa included in this study have columnar palisade cells that are extremely narrow and packed closely together. This layer allows little penetration of light, but exposure of the leaf undersurface may provide illumination of spongy mesophyll chloroplasts in these plants.
[23]	BJ?RKMAN O, HOMLMGREN P. Adaptability of the photosynthetic apparatus to light intensity in ecotypes from exposed and shaded habitats .Physoil Plantarum, 1963,16( 4) : 889-914.https://doi.org/10.1111/j.1399-3054.1963.tb08366.x URL [本文引用: 1]摘要 First page of article
[24]	秦玉芝, 邢铮, 邹剑锋, 何长征, 李炎林, 熊兴耀. 持续弱光胁迫对马铃薯苗期生长和光合特性的影响 . 中国农业科学, 2014, 47(3): 537-545.https://doi.org/10.3864/j.issn.0578-1752.2014.03.013 URL Magsci [本文引用: 1]摘要 1College of Horticulture and Landscape, Hunan Agricultural University, Changsha 410128; 2Center of Analytical Service, Hunan Agricultural University, Changsha 410128; 3Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences, Beijing 100081) Abstract: 【Objective】 The aim of this study was to investigate the effects of sustained weak light on growing conditions and photosynthetic characteristics of potato leaves at the seedling stage and to offer a theoretical basis for screening and utilization of new resistant potato resources to weak light.【Method】 The original cultivated potato Yan (S. tuberosum subsp. andigena var. yanacochense) and cultivated potato Favorita (S. tuberosum) were grown in a growth chamber at 50 μmol61m-261s-1 and 350 μmol61m-261s-1 light, respectively, using potato tuber (50 g) growing in medium. After 1 month treatment, the growth of the test materials was measured. Plant phenotype, the chlorophyll content, net photosynthetic rate, photosynthetic curve, the CO2 response curve were tested by LI-COR 6400 portable photosynthesis system. The stomatal morphological characters were measured using JEM-1200EX scanning electron micrography. The ultrastructure of chloroplast was observed by transmission electron microscope using the blade section after uranium acetate-citrate double staining.【Result】There was a difference between the treatment and the control in leaf size, morphology of plant. Favorita had weak vigour, slender seedling with smaller pale green foliage. Foliage and branch induction of Yan were very difficult and grew extremely slowly in or after treatment. Sustained light treatment significantly reduced the apparent quantum yield (AQE), light saturation point (LSP), maximum net photosynthetic rate (Pnmax), CO2 saturation concentration (CSP), and the chlorophyll content efficiency, and improved apparent carboxylation rate (EC), and CO2 compensation concentration (CCP) simultaneously. The light compensation point (LCP) of Yan was activated in the treatment. While the LCP and EC of Favorita were reverse. The EC, CSP of Yan had no difference between the treatment and the control. Growing under weak light, the stomatal density, chloroplast number declined. The numbers of grana and grana lamellae increased in Favorita. On the contrary, the number of grana lamellae decreased in Yan. 【Conclusion】 Foliage differentiation of potato was hypersensitivity to light. There was a significant genotypic difference. The development and growth of the sensitive genotypes were hindered when plants acclimated to low light (50 μmol61m-261s-1). The damage could not be resumed by restoring strong light later. The sustained weak light treatment at seedling stage depressed photosynthetic rate (Pn) and dark respiration rate (Rday). Saturation point of plant fell off along with the light intensities reduced. Following the dark respiration rate (Rday) being decreased, Favorita potato could adapt to lower light intensity by reducing the light compensation point. On the contrary, Yan could not adapt to lower light intensity. Appeared organic matter accumulated difficulties because of the improved light compensation point (LCP), the narrowing effective intensity range and the increased dark respiration rate (Rday). Under sustained weak light stress, the mesophyll cell arrangement was changed , and the stomatal density, the stomatal apparatus and numbers of chloroplasts decreased significantly, the stomatal apparatus length-width ratio had a tendency to increase, and chlorophyll composition ratio changed. Favorita potato improved the capture ability of effective light sources by increasing the number of chloroplast grana, grana lamella and chlorophyll b. Yan potato had subdued capturing ability of effective light and CO2 affinity because the formation of chloroplast grana was affected and the number of chloroplast grana, grana lamella and stomatal density decreased. QIN Y Z, XING Z, ZOU J F, HE C Z, LI Y L, XIONG X Y.Effects of sustained weak light on seedling growth and photosynthetic characteristics of potato seedlings .Scientia Agricultura Sinica, 2014, 47(3): 537-545. (in Chinese)https://doi.org/10.3864/j.issn.0578-1752.2014.03.013 URL Magsci [本文引用: 1]摘要 1College of Horticulture and Landscape, Hunan Agricultural University, Changsha 410128; 2Center of Analytical Service, Hunan Agricultural University, Changsha 410128; 3Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences, Beijing 100081) Abstract: 【Objective】 The aim of this study was to investigate the effects of sustained weak light on growing conditions and photosynthetic characteristics of potato leaves at the seedling stage and to offer a theoretical basis for screening and utilization of new resistant potato resources to weak light.【Method】 The original cultivated potato Yan (S. tuberosum subsp. andigena var. yanacochense) and cultivated potato Favorita (S. tuberosum) were grown in a growth chamber at 50 μmol61m-261s-1 and 350 μmol61m-261s-1 light, respectively, using potato tuber (50 g) growing in medium. After 1 month treatment, the growth of the test materials was measured. Plant phenotype, the chlorophyll content, net photosynthetic rate, photosynthetic curve, the CO2 response curve were tested by LI-COR 6400 portable photosynthesis system. The stomatal morphological characters were measured using JEM-1200EX scanning electron micrography. The ultrastructure of chloroplast was observed by transmission electron microscope using the blade section after uranium acetate-citrate double staining.【Result】There was a difference between the treatment and the control in leaf size, morphology of plant. Favorita had weak vigour, slender seedling with smaller pale green foliage. Foliage and branch induction of Yan were very difficult and grew extremely slowly in or after treatment. Sustained light treatment significantly reduced the apparent quantum yield (AQE), light saturation point (LSP), maximum net photosynthetic rate (Pnmax), CO2 saturation concentration (CSP), and the chlorophyll content efficiency, and improved apparent carboxylation rate (EC), and CO2 compensation concentration (CCP) simultaneously. The light compensation point (LCP) of Yan was activated in the treatment. While the LCP and EC of Favorita were reverse. The EC, CSP of Yan had no difference between the treatment and the control. Growing under weak light, the stomatal density, chloroplast number declined. The numbers of grana and grana lamellae increased in Favorita. On the contrary, the number of grana lamellae decreased in Yan. 【Conclusion】 Foliage differentiation of potato was hypersensitivity to light. There was a significant genotypic difference. The development and growth of the sensitive genotypes were hindered when plants acclimated to low light (50 μmol61m-261s-1). The damage could not be resumed by restoring strong light later. The sustained weak light treatment at seedling stage depressed photosynthetic rate (Pn) and dark respiration rate (Rday). Saturation point of plant fell off along with the light intensities reduced. Following the dark respiration rate (Rday) being decreased, Favorita potato could adapt to lower light intensity by reducing the light compensation point. On the contrary, Yan could not adapt to lower light intensity. Appeared organic matter accumulated difficulties because of the improved light compensation point (LCP), the narrowing effective intensity range and the increased dark respiration rate (Rday). Under sustained weak light stress, the mesophyll cell arrangement was changed , and the stomatal density, the stomatal apparatus and numbers of chloroplasts decreased significantly, the stomatal apparatus length-width ratio had a tendency to increase, and chlorophyll composition ratio changed. Favorita potato improved the capture ability of effective light sources by increasing the number of chloroplast grana, grana lamella and chlorophyll b. Yan potato had subdued capturing ability of effective light and CO2 affinity because the formation of chloroplast grana was affected and the number of chloroplast grana, grana lamella and stomatal density decreased.
[25]	CAMPBELL C D, SAGE R F, KOCACINAR F, WAY D A.Estimation of the whole plant CO₂ compensation point of tobacco ( Nicotiana tabacum L.). Global Change Biology, 2010, 11: 1956-1967. [本文引用: 1]
[26]	王志强, 何方, 牛良, 刘淑娥, 宗学普, 何方. CO₂施肥对大棚油桃光合作用及产量品质的影响 . 果树学报, 2001, 18(2): 75-79.https://doi.org/10.3969/j.issn.1009-9980.2001.02.004 URL [本文引用: 1]摘要棚内CO2 浓度日变化幅度远大于棚外 ;在天气晴好、光合旺盛、通风受阻时 ,棚内CO2 匮乏可成为光合作用的主要限制因子。在保护地条件下 ,油桃光合速率 (Pn)日变化由露地的双峰曲线变为三峰曲线 ,最大Pn比露地提前 2h ,在 8时左右出现 ;光合“午休”现象不明显 ;晴天棚内日平均Pn为 5 .97(CO2 ) μmol/m2 ·s,比露地降低17.2 5 %。棚内CO2 加富对油桃光合作用和产量、品质有重要影响 ,较大幅度地提高了上午 8时～ 12时之间的Pn和光能利用率 ,日平均Pn 6 .95 (CO2 ) μmol/m2 ·s ,比对照提高 2 5 .90 %。树体生长健壮 ,生物量 (未含果实 )增加 12 .4% ,比叶重增大 ,产量提高 19.80 % ,品质改善 WANG Z Q, HE F, NIU L A, LIU SH E, ZONG X P, HE F.The effects of CO₂ enrichment on photosynthesis, yield and quality of nectarine in greenhouse .Journal of Fruit Science, 2001, 18(2):75-79. (in Chinese)https://doi.org/10.3969/j.issn.1009-9980.2001.02.004 URL [本文引用: 1]摘要棚内CO2 浓度日变化幅度远大于棚外 ;在天气晴好、光合旺盛、通风受阻时 ,棚内CO2 匮乏可成为光合作用的主要限制因子。在保护地条件下 ,油桃光合速率 (Pn)日变化由露地的双峰曲线变为三峰曲线 ,最大Pn比露地提前 2h ,在 8时左右出现 ;光合“午休”现象不明显 ;晴天棚内日平均Pn为 5 .97(CO2 ) μmol/m2 ·s,比露地降低17.2 5 %。棚内CO2 加富对油桃光合作用和产量、品质有重要影响 ,较大幅度地提高了上午 8时～ 12时之间的Pn和光能利用率 ,日平均Pn 6 .95 (CO2 ) μmol/m2 ·s ,比对照提高 2 5 .90 %。树体生长健壮 ,生物量 (未含果实 )增加 12 .4% ,比叶重增大 ,产量提高 19.80 % ,品质改善
[27]	张艳丽, 张重义, 李友军, 刘辉, 李娟. 不同密度下牛膝叶绿素荧光特性比较 . 生态学杂志, 2008, 27(7): 1089-1094.URL Magsci [本文引用: 1]摘要 <P><FONT face=Verdana>在大田条件下，测定了不同密度下（A： 500 000；B： 330 000；C： 250 000；</FONT></P><P><FONT face=Verdana>D</FONT><FONT face=Verdana>：200 000株·hm<SUP>-2</SUP>）牛膝（<EM>Achyranthes bidentata</EM>）的叶面积动态和倒二叶叶绿素含量、叶绿素荧光动力学参数以及根产量和牛膝多糖含量，了解了不同密度下牛膝的叶绿素荧光特性，为牛膝规范化种植提供理论依据。结果表明: 不同密度下叶面积指数和叶绿素含量最高值均在苗后45 d，分别为A>B>C>D和B>C>A>D；光系统Ⅱ中的<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>o</SUB>、<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>m</SUB>、<EM>Ф</EM><SUB>PSⅡ</SUB>、<EM>q</EM><SUB>P</SUB>、<EM>ETR</EM>均是先上升后下降，<EM>q</EM>N则升高；同一生育时期B处理<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>o</SUB>、<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>m</SUB>、<EM>Ф</EM><SUB>PSⅡ</SUB>、<EM>q</EM>N和<EM>ETR</EM>则表现为最高，且B处理受强光抑制的程度最小，B处理表现较高的光合作用；根产量和牛膝多糖含量与<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>o</SUB>、<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>m</SUB>呈正相关，与<EM>q</EM>N呈负相关；在出穗55 d，<EM>q</EM>N与多糖含量达显著负相关；B处理为最适密度。</FONT></P> ZHANG Y L, ZHANG Z Y, Li Y J, LIU H, L J. Chlorophyll fluorescence characteristics of Achyranthes bidentata under different planting densities. Chinese Journal of Ecology, 2008, 27(7): 1089-1094. (in Chinese)URL Magsci [本文引用: 1]摘要 <P><FONT face=Verdana>在大田条件下，测定了不同密度下（A： 500 000；B： 330 000；C： 250 000；</FONT></P><P><FONT face=Verdana>D</FONT><FONT face=Verdana>：200 000株·hm<SUP>-2</SUP>）牛膝（<EM>Achyranthes bidentata</EM>）的叶面积动态和倒二叶叶绿素含量、叶绿素荧光动力学参数以及根产量和牛膝多糖含量，了解了不同密度下牛膝的叶绿素荧光特性，为牛膝规范化种植提供理论依据。结果表明: 不同密度下叶面积指数和叶绿素含量最高值均在苗后45 d，分别为A>B>C>D和B>C>A>D；光系统Ⅱ中的<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>o</SUB>、<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>m</SUB>、<EM>Ф</EM><SUB>PSⅡ</SUB>、<EM>q</EM><SUB>P</SUB>、<EM>ETR</EM>均是先上升后下降，<EM>q</EM>N则升高；同一生育时期B处理<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>o</SUB>、<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>m</SUB>、<EM>Ф</EM><SUB>PSⅡ</SUB>、<EM>q</EM>N和<EM>ETR</EM>则表现为最高，且B处理受强光抑制的程度最小，B处理表现较高的光合作用；根产量和牛膝多糖含量与<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>o</SUB>、<EM>F</EM><SUB>v</SUB>/<EM>F</EM><SUB>m</SUB>呈正相关，与<EM>q</EM>N呈负相关；在出穗55 d，<EM>q</EM>N与多糖含量达显著负相关；B处理为最适密度。</FONT></P>
[28]	罗青红, 李志军, 伍维模, 韩路. 胡杨、灰叶胡杨光合及叶绿素荧光特性的比较研究 . 西北植物学报, 2006, 26(5):983-988.https://doi.org/10.3321/j.issn:1000-4025.2006.05.018 URL [本文引用: 1]摘要比较研究了两年生胡杨和灰叶胡杨叶片的光合及叶绿素荧光特性.结果表明：胡杨和灰叶胡杨均表现出净光合速率（Pn）日变化呈单峰曲线,只是在12：00时胡杨的Pn略微降低;气孔导度日变化均呈单峰型曲线;胞间CO 2浓度日变化呈近“V”字型曲线.相同的试验条件下,胡杨的净光合速率、气孔导度高于灰叶胡杨,净光合速率与气孔导度峰值出现在上午10：00时;胡杨胞间CO2浓度低于灰叶胡杨,胞间CO2浓度最低值均出现在中午14：00时.经充分暗适应后的叶片叶绿素荧光参数初始荧光、PSⅡ原初光能转换效率和PS Ⅱ潜在活性均为胡杨显著大于灰叶胡杨.以太阳光为光化学光,测定的叶绿素荧光参数日变化显示,胡杨PSⅡ实际的光化学反应量子效率、非循环电子传递速率、光化学猝灭系数均大于灰叶胡杨,而非光化学猝灭系数却较灰叶胡杨小.胡杨与灰叶胡杨在光合与叶绿素荧光特性上的差异,是胡杨更能适应干旱荒漠区高光、高温与低相对空气湿度环境,从而表现出高净光合速率的部分生理学原因之一. LUO Q H, LI Z J, WU W M, HAN L.Comparative study of photosynthetic and chlorophyll fluorescence characteristics of Populus euphratica and P. pruinosa. Acta Botanica Boreali-Occidentalia Sinica, 2006, 26(5): 983-988. (in Chinese)https://doi.org/10.3321/j.issn:1000-4025.2006.05.018 URL [本文引用: 1]摘要比较研究了两年生胡杨和灰叶胡杨叶片的光合及叶绿素荧光特性.结果表明：胡杨和灰叶胡杨均表现出净光合速率（Pn）日变化呈单峰曲线,只是在12：00时胡杨的Pn略微降低;气孔导度日变化均呈单峰型曲线;胞间CO 2浓度日变化呈近“V”字型曲线.相同的试验条件下,胡杨的净光合速率、气孔导度高于灰叶胡杨,净光合速率与气孔导度峰值出现在上午10：00时;胡杨胞间CO2浓度低于灰叶胡杨,胞间CO2浓度最低值均出现在中午14：00时.经充分暗适应后的叶片叶绿素荧光参数初始荧光、PSⅡ原初光能转换效率和PS Ⅱ潜在活性均为胡杨显著大于灰叶胡杨.以太阳光为光化学光,测定的叶绿素荧光参数日变化显示,胡杨PSⅡ实际的光化学反应量子效率、非循环电子传递速率、光化学猝灭系数均大于灰叶胡杨,而非光化学猝灭系数却较灰叶胡杨小.胡杨与灰叶胡杨在光合与叶绿素荧光特性上的差异,是胡杨更能适应干旱荒漠区高光、高温与低相对空气湿度环境,从而表现出高净光合速率的部分生理学原因之一.
[29]	简在友, 王文全, 孟丽, 许桂芳,王秋玲, 李卫东, 俞敬波. 芍药组内不同类群间光合特性及叶绿素荧光特性比较 . 植物生态学报, 2010, 34(12): 1463-1471.https://doi.org/10.3773/j.issn.1005-264x.2010.12.013 URL Magsci [本文引用: 1]摘要为了比较分析芍药组不同类群光合生理差异以及它们对各自不同光照环境的适应性,测定了芍药组(Paeonia sect.Paeonia)4种2变种的光合日变化、光响应曲线、CO2响应曲线,以及叶绿素荧光特性.结果显示,芍药组中不同类群之间光合速率差异明显,各个种类的"午休"程度不同,芍药(Paeonia lactiflora)和毛果芍药(P.lactiflora var.trichocarpa)的强光抑制现象没有川赤芍(P.veitchii)、美丽芍药(P.mairei)和窄叶芍药(P.anomata)明显.叶绿素荧光特征能够反映芍药组不同类群光合生理的差异.芍药组内不同类群地理分布的差异能部分从光合生理特征的适应性方面得到解释. JIAN Z Y, WANG W Q, MENG L, XU G F, WANG Q L, LI W D, YU J B.Comparison of photosynthetic and fluorescence characteristics among taxa in Paeonia sect. Paeonia .Chinese Journal of Plant Ecology, 2010, 34(12): 1463-1471. (in Chinese)https://doi.org/10.3773/j.issn.1005-264x.2010.12.013 URL Magsci [本文引用: 1]摘要为了比较分析芍药组不同类群光合生理差异以及它们对各自不同光照环境的适应性,测定了芍药组(Paeonia sect.Paeonia)4种2变种的光合日变化、光响应曲线、CO2响应曲线,以及叶绿素荧光特性.结果显示,芍药组中不同类群之间光合速率差异明显,各个种类的"午休"程度不同,芍药(Paeonia lactiflora)和毛果芍药(P.lactiflora var.trichocarpa)的强光抑制现象没有川赤芍(P.veitchii)、美丽芍药(P.mairei)和窄叶芍药(P.anomata)明显.叶绿素荧光特征能够反映芍药组不同类群光合生理的差异.芍药组内不同类群地理分布的差异能部分从光合生理特征的适应性方面得到解释.
[30]	吴飞燕,伊力塔,李修鹏,殷秀敏, 刘美华, 余树全.不同光照强度对石栎幼苗叶绿素含量及叶绿素荧光参数的影响 . 东北农业大学学报,2012, 43(4): 88-92.https://doi.org/10.3969/j.issn.1005-9369.2012.04.016 URL [本文引用: 1]摘要以亚热带常绿阔叶林中的常见种石栎(Lithocarpus glaber)为研究材料,测定了不同光强条件(100%、75%和50%自然光强)下幼苗叶绿素荧光参数及叶绿素含量,探讨了石栎生长对光强的适应性。结果表明,随着光照强度的减弱,石栎叶片的叶绿素含量上升,3种光强条件下叶片光系统Ⅱ的实际光化学效率(ΦPSII)、最大光能转化效率(Fv/Fm)和潜在活性(Fv/Fo)的季节变化规律相似,高光强下的ΦPSII、Fv/Fm和Fv/Fo值相对较小,说明石栎幼苗能适应较大的光强幅度,同时对低、弱光强有一定的抗逆性。 WU F Y, YI L T, LI X P, YIN X M, LIU M H, YU SH Q.Effect of different light intensity on intensity chlorophyll content and chlorophyll fluorescence in Lithocarpus glaber. Journal of Northeast Agricultural University, 2012, 43(4): 88-92. (in Chinese)https://doi.org/10.3969/j.issn.1005-9369.2012.04.016 URL [本文引用: 1]摘要以亚热带常绿阔叶林中的常见种石栎(Lithocarpus glaber)为研究材料,测定了不同光强条件(100%、75%和50%自然光强)下幼苗叶绿素荧光参数及叶绿素含量,探讨了石栎生长对光强的适应性。结果表明,随着光照强度的减弱,石栎叶片的叶绿素含量上升,3种光强条件下叶片光系统Ⅱ的实际光化学效率(ΦPSII)、最大光能转化效率(Fv/Fm)和潜在活性(Fv/Fo)的季节变化规律相似,高光强下的ΦPSII、Fv/Fm和Fv/Fo值相对较小,说明石栎幼苗能适应较大的光强幅度,同时对低、弱光强有一定的抗逆性。
[31]	VAN KOOTEN O, SNEL J F H.Photosynthesis Research , 1990, 25: 147-150. [本文引用: 1]
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[33]	STEFANOS K, IONNIST T, ELEFTHERIA Z, NIKOLAOS N.Rootstock effects on the adaptive strategies of grapevine (Vitis vinifera L. cv. Cabernet-Sauvignon) under contrasting water status: Leaf physiological and structural responses. Agriculture, Ecosystems & Environment, 2008, 128(1): 86-96.https://doi.org/10.1016/j.agee.2008.05.006 URL [本文引用: 1]摘要 The effects of two rootstocks (1103P and SO4) on water status, gas exchange, leaf structural characteristics and vine growth were studied in a factorial experiment over 2 years (2005–2006), in field-grown grapevines of cv. Cabernet–Sauvignon ( Vitis vinifera L.), subjected to three irrigation levels (FI: 100% of evapotranspiration, DI: 50% of evapotranspiration and NI: not irrigated). The experiment was conducted on 10-year-old vines, grown under the semi-arid conditions of central Greece. There was a significant depressive effect in water status (estimated by midday stem water potential, Ψ s) and net assimilation rate ( A) of Cabernet–Sauvignon with decreasing water supply, on both rootstocks. The down regulation of A was mainly due to decreases in stomatal conductance ( g s). NI treatment showed significantly higher intrinsic water use efficiency ( A/ g s) and lower leaf carbon isotope discrimination ( Δ) compared to irrigated treatments. Leaf area index (LAI) and pruning weight (PW) were found to be higher under FI treatment but this effect was more pronounced on the 1103P-grafted vines. Canopy growth was strongly controlled by rootstock, being significantly higher on 1103P, presumably due to its higher capacity to access soil water. Rootstock genotype affected scion water status via its effect on whole-vine transpiration. Cabernet–Sauvignon photosynthesis response to water status was not altered by rootstock. The only apparent difference between rootstocks was the higher A of SO4-grafted vines under NI treatment, associated with higher Ψ s. A/ g s and Δ were only altered under FI treatment, with 1103P showing more efficient use of available soil water than SO4. Vines on SO4 had a lower specific leaf area (SLA) as a result of lower growth rate than 1103P, which possibly provided a supplemental mechanism to maintain leaf water status and gas exchange. Concluding, SO4 could be better adapted on fertile soils under non-limiting water supply due to its capacity to achieve a balanced vegetative and reproductive growth while 1103P, a more water efficient rootstock, would be better to grow in semi-arid regions where water limitation occurs.
[34]	郑秋玲, 王慧, 韩真, 翟衡. 赤霞珠不同砧穗组合对中后期光合能力及果实品质的影响 . 中外葡萄与葡萄酒, 2010(3): 10-13.https://doi.org/10.3969/j.issn.1004-7360.2010.05.002 URL [本文引用: 1]摘要为了探讨砧木和品系对葡萄生长发育的影响，以赤霞珠葡萄的不同砧穗组合为试材，测定其生长季中后期光合荧光参数、果实品质及休眠期枝条和根系贮藏营养。结果表明：相同品系不同砧木，CS—169／SO4组合中期（8月份）光合速率最高，而CS-169／1103P组合后期（9、10月份）光合速率最高；CS-169／1103P组合的Fv／Fo、Fm、ETR、ФPSⅡ指标在不同时期均处于最高水平；同一砧木不同品系光合和荧光参数均以CS-169品系优于R5、FV5品系；以1103P为砧木的果实糖含量高于SO4砧，不同品系CS-169〉CS．FV5〉CS—R5，组合CS-169／SO4果实酸含量最高；枝条和根系的贮藏C、N营养以CS-169／1103P组合含量最高。综合看，砧木的影响大于品系，赤霞珠以1103P为砧木的各砧穗组合在光合荧光特性、果实品质、树体贮藏营养方面优于以SO4为砧木的组合。 ZHENG Q L, WANG H, HAN Z, ZHAI H.The influences of several scion-cultivar/rootstock combinations on photosynthesis during middle and late growing seasons and berry quality of Cabernet Sauvignon grapevine .Sino-Overseas Grapevine & Wine, 2010(3): 10-13. (in Chinese)https://doi.org/10.3969/j.issn.1004-7360.2010.05.002 URL [本文引用: 1]摘要为了探讨砧木和品系对葡萄生长发育的影响，以赤霞珠葡萄的不同砧穗组合为试材，测定其生长季中后期光合荧光参数、果实品质及休眠期枝条和根系贮藏营养。结果表明：相同品系不同砧木，CS—169／SO4组合中期（8月份）光合速率最高，而CS-169／1103P组合后期（9、10月份）光合速率最高；CS-169／1103P组合的Fv／Fo、Fm、ETR、ФPSⅡ指标在不同时期均处于最高水平；同一砧木不同品系光合和荧光参数均以CS-169品系优于R5、FV5品系；以1103P为砧木的果实糖含量高于SO4砧，不同品系CS-169〉CS．FV5〉CS—R5，组合CS-169／SO4果实酸含量最高；枝条和根系的贮藏C、N营养以CS-169／1103P组合含量最高。综合看，砧木的影响大于品系，赤霞珠以1103P为砧木的各砧穗组合在光合荧光特性、果实品质、树体贮藏营养方面优于以SO4为砧木的组合。
[35]	陈继峰. 葡萄砧木品种的研究现状与展望 . 果树学报, 2000, 17(2): 138-146.https://doi.org/10.3969/j.issn.1009-9980.2000.02.013 URL [本文引用: 1]摘要在大量文献的基础上，主要描述了用作砧木和砧木育种材料的葡萄重要的种和世界各国广泛应用的砧木品种的主要特性；综述了一些砧木的抗病虫、耐寒、耐旱、抗湿等性能；砧木对所嫁接品种的影响和砧木对矿物质的吸收利用以及嫁接亲和性等方面的研究成果与进展；对我国砧木工作的研究提出了建议。 CHEN J F.The Status of Research on Grape Rootstock Varieties and Its Prospect .Journal of Fruit Science, 2000, 17(2): 138-146. (in Chinese)https://doi.org/10.3969/j.issn.1009-9980.2000.02.013 URL [本文引用: 1]摘要在大量文献的基础上，主要描述了用作砧木和砧木育种材料的葡萄重要的种和世界各国广泛应用的砧木品种的主要特性；综述了一些砧木的抗病虫、耐寒、耐旱、抗湿等性能；砧木对所嫁接品种的影响和砧木对矿物质的吸收利用以及嫁接亲和性等方面的研究成果与进展；对我国砧木工作的研究提出了建议。
[36]	王海波, 王孝娣, 史祥宾, 王宝亮, 郑晓翠, 刘凤之. 葡萄不同品种对设施环境的适应性 . 中国农业科学, 2013, 46(6): 1213-1220.https://doi.org/10.3864/j.issn.0578-1752.2013.06.015 URL Magsci [本文引用: 1]摘要【目的】选择适宜设施栽培环境的葡萄品种。【方法】以22个5年生设施葡萄常用品种为试验材料，研究不同葡萄品种对设施环境的适应性，明确影响设施葡萄连年丰产的重要环境因子。【结果】设施栽培环境条件下连年丰产能力强的品种其耐弱光能力强，但耐低浓度CO2和耐高温能力不一定强；弱光是影响设施葡萄连年丰产的主要环境因素，而低浓度CO2和高温不是影响设施葡萄连年丰产的主要环境因素。【结论】红旗特早玫瑰、紫珍香、无核白鸡心、无核早红、红标无核、87-1、乍娜、莎巴珍珠、奥迪亚无核、香妃和红香妃等品种对设施栽培环境具有较强的适应性，连年丰产能力强。弱光是影响设施葡萄连年丰产的重要环境因素。 WANG H B, WANG X D, SHI X B, WANG B L, ZHENG X C, LIU F Z.Environmental adaptability of different grape cultivars in greenhouse .Scientia Agricultura Sinica, 2013, 46(6): 1213-1220. (in Chinese)https://doi.org/10.3864/j.issn.0578-1752.2013.06.015 URL Magsci [本文引用: 1]摘要【目的】选择适宜设施栽培环境的葡萄品种。【方法】以22个5年生设施葡萄常用品种为试验材料，研究不同葡萄品种对设施环境的适应性，明确影响设施葡萄连年丰产的重要环境因子。【结果】设施栽培环境条件下连年丰产能力强的品种其耐弱光能力强，但耐低浓度CO2和耐高温能力不一定强；弱光是影响设施葡萄连年丰产的主要环境因素，而低浓度CO2和高温不是影响设施葡萄连年丰产的主要环境因素。【结论】红旗特早玫瑰、紫珍香、无核白鸡心、无核早红、红标无核、87-1、乍娜、莎巴珍珠、奥迪亚无核、香妃和红香妃等品种对设施栽培环境具有较强的适应性，连年丰产能力强。弱光是影响设施葡萄连年丰产的重要环境因素。