谢辉^1,,
张雯^1,,,
韩守安¹,
王敏¹,
阿力木江·奥布力²,
潘明启¹,
艾尔买克·才卡斯木¹,
张平³
1.新疆农业科学院园艺作物研究所/农业部新疆地区果树科学观测试验站 乌鲁木齐 830091
2.和田地区林业和草原局 和田 848000
3.新疆农业科学院农产品贮藏加工研究所 乌鲁木齐 830091
基金项目: 国家自然科学基金地区基金项目31560138
新疆维吾尔自治区优秀青年科技人才培养项目qn2016yx0234

详细信息

作者简介:谢辉, 研究方向为果园生态与果品加工。E-mail: xhxjnky@163.com

通讯作者:张雯, 研究方向为果园生态与果实品质调控。E-mail: zwxilin@126.com

中图分类号:S512.1;S311

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出版历程

收稿日期:2020-10-22
录用日期:2021-01-19
刊出日期:2021-04-01

Effect of shading degree on the grain yield and photosynthetic characteristics of wheat at the grain filling stage in an almond-winter wheat intercropping system

XIE Hui^1,,
ZHANG Wen^1,,,
HAN Shou'an¹,
WANG Min¹,
Alimujiang Aubrey²,
PAN Mingqi¹,
Aiermaike Caikasimu¹,
ZHANG Ping³
1. Institute of Horticultural Crops, Xinjiang Academy of Agricultural Sciences/Scientific Observing and Experimental Station of Pomology(Xinjiang), Urumqi 830091, China
2. Forestry and Grassland Administration of Hetian Prefecture, Hetian 848000, China
3. Institute of Agricultural Product Storage and Processing, Xinjiang Academy of Agricultural Sciences, Urumqi 830091, China
Funds: the National Natural Science Foundation of China31560138
the Outstanding Young Science and Technology Talent Training Project of Xinjiang Uygur Autonomous Regionqn2016yx0234

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Corresponding author:ZHANG Wen, E-mail: zwxilin@126.com

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摘要
摘要:为探明果粮间作模式下树冠遮阴对间作冬小麦灌浆期光合特性和产量的影响，为新疆南疆果粮间作模式的选择和优化提供理论依据，本研究以扁桃-冬小麦间作模式为研究对象，设置重度遮阴和轻度遮阴2个处理，以单作小麦为对照，对间作区域光合有效辐射（PAR）、小麦产量及灌浆期叶片光合色素和可溶性蛋白含量、光响应曲线、荧光动力学参数、群体光合速率进行测定。结果表明：间作区域遮阴程度受树冠结构和间作区域距树体距离共同影响，主干分层形扁桃（重度遮阴）近冠和远冠间作区域小麦灌浆期PAR日均值为自然光强的18.61%和25.90%，小冠圆头形扁桃（轻度遮阴）对应值为56.00%和64.53%。与扁桃间作导致小麦旗叶Chla+b含量和Chla/b比值降低；PAR日均值≤56.00%自然光时，小麦旗叶可溶性蛋白质含量显著降低；PAR日均值≤25.90%自然光时，旗叶光系统Ⅱ的实际光合效率（ΦSPⅡ）、光化学淬灭系数（qP）、最大净光合速率（P_max）及群体光合速率日均值均显著降低；PAR日均值≥64.53%自然光时小麦群体光合速率存在明显的补偿现象。综上，扁桃-冬小麦间作模式下，小麦光合能力及产量与遮阴程度密切相关，重度遮阴导致小麦单叶和群体光合能力及产量显著降低；轻度遮阴条件下，小麦旗叶光合能力无显著变化，群体光合速率存在明显的补偿现象，对产量无显著影响。
关键词:扁桃-冬小麦间作/
树形/
遮阴强度/
间作区域/
光合特性/
产量
Abstract:Fruit tree-wheat intercropping is practiced in large parts of southern Xinjiang, a region where agroforestry intercropping is the main type of agricultural production. In the present study, a field experiment was conducted to investigate the effects of the shading degree from fruit tree canopies on the grain yield and photosynthetic characteristics of wheat at the grain filling stage. This study aimed to provide information for the selection of management standards and the optimization of the intercropping system in southern Xinjiang. In the experiment, 'Xindong 20' (Triticum aestivum L. var. Xindong 20), the main winter wheat cultivar in South Xinjiang, was used as the research object, and two treatments (heavy and light shading) were established by cutting the almond (Amygdalus communis L.) canopy of the almond-winter wheat intercropping system; delayed open-central canopy (DC) and a semicircle small-canopy (SC), respectively. Monocultured wheat was used as the control. The yield and yield components, photosynthetic pigments and soluble protein contents, light response curves, chlorophyll fluorescence induction of the flag leaves, and the canopy apparent photosynthetic rate of the intercropped wheat were investigated in three areas: near the canopy west of the almond trees, near the canopy east of the almond trees, and far from the canopy. The results indicated that the shading degree of the tree canopy was closely associated with the canopy size and the distance between the intercropping area and the tree. The daily mean values of photosynthetically active radiation (PAR) in the intercropping areas near and far from the almond canopy were 18.61% and 25.90% with DC, and 56.00% and 64.53% with SC of the natural light intensity during the wheat-filling period, respectively. The content of Chla+b and the Chla/b ratio in the wheat flag leaves were reduced in both shading treatments. When the daily mean strength of PAR was ≤56.00% of the natural light intensity, the soluble protein content in the wheat flag leaves was significantly reduced. When the daily average intensity of PAR was reduced to 25.90% and 18.61% of the natural light, the indices of optical system Ⅱ actual photosynthetic efficiency (ΦSPⅡ), photochemical quenching coefficient (qP), and maximum net photosynthetic rate (P_max) of the flag leaves, as well as the daily mean canopy photosynthetic rate (CAP) value, were significantly reduced. Under the light shading intercropping condition, when the daily mean intensity of PAR reached ≥ 64.53% of the natural light intensity, the CAP value exhibited an obvious increased (compensation phenomenon). In the almond-winter wheat intercropping system, the influence of a weak light environment caused by the tree canopy on wheat photosynthetic capacity and yield was closely related to the degree of shading. Heavy shading resulted in a significant decrease in single-leaf P_n and CAP and caused a significant decrease in yield. Under light shading conditions, the photosynthetic capacity of wheat flag leaves did not change, while CAP had a significant compensation phenomenon and no significant influence on grain yield.
Key words:Almond-winter wheat crop intercropping/
Tree form/
Shading degree/
Intercropping area/
Photosynthetic characteristics/
Grain yield

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图1主干分层形重度遮阴(DC)和小冠形轻度遮阴(SC)扁桃树形结构示意图
Figure1.Sketch maps of delayed open-central canopy (DC, heavy shading) and semicircle small-canopy (SC, light shading) of almond


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图2扁桃-冬小麦间作模式及测点位置示意图
Figure2.Sketch map of almond-winter wheat intercropping system and positions of measuring points


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图3扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域小麦灌浆期光合有效辐射(PAR)的日变化
CK为单作; W-Zone1为树行西侧近冠区域, E-Zone1为树行东侧近冠区域, Zone2为西侧远冠区域。
Figure3.Diurnal variation trends of photosynthetic active radiation (PAR) at grain-filling stage of wheat in different intercropping areas under different canopy-shading degrees (different canopy structures of almond) of almond-winter wheat intercropping system
CK is the monoculture wheat. W-Zone1 is the intercropping area near canopy on the west side of tree line, E-Zone1 is the intercropping area near canopy on the east side of tree line, Zone2 is the intercropping area far from tree canopy on the west side of tree line.


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图4扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域冬小麦灌浆期旗叶光响应曲线
CK为单作小麦, Zone1为近冠区, Zone2为远冠区。
Figure4.Light response curves of net photosynthetic rate (P_n) of flag leaf of winter wheat at filling stage in different intercropping areas under different canopy-shading degrees of almond-winter wheat intercropping system
CK is the monocultured winter wheat. Zone1 is the area near canopy, Zone2 is the area far from canopy.


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图5扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域小麦灌浆期群体光合速率(CAP)的日变化
CK为单作对照; DC: 主干分层形重度遮阴; SC: 小冠形轻度遮阴; W-Zone1为树行西侧近冠区域, E-Zone1为树行东侧近冠区域, Zone2为西侧远冠区域。不同小写字母表示对照(CK)及不同遮阴强度及间作区域间在P < 0.05水平差异显著。
Figure5.Daily dynamics of canopy apparent photosynthetic rate (CAP) of winter wheat population in different intercropping areas under different canopy-shading degrees of almond-winter wheat intercropping system
CK is the monocultued wheat. DC is the treatment of heavy shading of delayed open-central canopy; SC is the treatment of light shading of semicircle small-canopy. W-Zone1 is the intercropping area near canopy on the west side of tree line, E-Zone1 is the intercropping area near canopy on the east side of tree line, Zone2 is the intercropping area far from tree canopy on the west side of tree line. Different lowercase letters indicate significant difference among control (CK) and different intercropping areas under different shading degrees at P < 0.05.


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图6扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域小麦灌浆期群体呼吸速率(CRR)的日变化
CK为单作对照; DC: 主干分层形重度遮阴; SC: 小冠形轻度遮阴; W-Zone1为树行西侧近冠区域, E-Zone1为树行东侧近冠区域, Zone2为西侧远冠区域。不同小写字母表示对照(CK)及不同遮阴强度及间作区域间在P < 0.05水平差异显著。
Figure6.Daily dynamics of canopy apparent photosynthetic rate (CRR) of winter wheat population in different intercropping areas under different canopy-shading degrees of almond-winter wheat intercropping system
CK is the monocultued wheat. DC is the treatment of heavy shading of delayed open-central canopy; SC is the treatment of light shading of semicircle small-canopy. W-Zone1 is the intercropping area near canopy on the west side of tree line, E-Zone1 is the intercropping area near canopy on the east side of tree line, Zone2 is the intercropping area far from tree canopy on the west side of tree line. Different lowercase letters indicate significant difference among control (CK) and different intercropping areas under different shading degrees at P < 0.05.


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表1主干分层形重度遮阴(DC)和小冠形轻度遮阴(SC)扁桃盛果期树形结构指标
Table1.Canopy structure indexes of delayed open-central canopy (DC) and semicircle small-canopy (SC) of almond at full bearing period

树形结构 Canopy structure	单株主枝数 Number of main branch	一年生枝量 Number of annual branch	树冠体积 Crown volume (m3)	单位体积枝量 Number of shoot (shoots·m-3)	一年生枝数量区域分布比例 Annual branch distribution ratio (%)
					外围 Outside canopy	中部 Middle canopy	内堂 Inner canopy
DC	8.4	5590.0	55.96	99.89	60	30	10
SC	7.3	1276.8	20.36	62.70	60	40	0

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表2扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域冬小麦灌浆期叶片叶绿素和可溶性蛋白含量
Table2.Contents of chlorophyll and soluble protein in leaves of winter wheat at filling stage in different intercropping areas under different canopy-shading degrees of almond-winter wheat intercropping system

处理 Treatment	间作小麦区域 Area of intercropped wheat	叶绿素含量?Chlorophyll content (mg·g-1)			叶绿素 a/b Chla/b	可溶性蛋白含量 Soluble protein content (mg·g-1)
		叶绿素a Chl. a	叶绿素b Chl. b	叶绿素a+b ? Chl. a+b
轻度遮阴 Light shading	近冠区?Near canopy	2.25±0.21b	0.96±0.08a	3.21±0.27b	2.39±0.14b	14.04±0.76b
	远冠区?Far from canopy	2.42±0.22a	0.85±0.07b	3.27±0.28a	2.90±0.18a	18.44±1.36a
重度遮阴 Heavy shading	近冠区?Near canopy	2.03±0.17c	1.05±0.09a	3.08±0.24c	1.97±0.87c	11.73±0.89c
	远冠区?Far from canopy	1.98±0.12d	0.68±0.06c	2.66±0.21d	2.84±0.12d	12.46±1.03d
单作小麦?Monocultured wheat (CK)		2.52±0.23a	0.84±0.06b	3.36±0.28a	3.04±0.13a	17.06±1.14a
表内数据为平均值±标准误, 同列数据后不同小写字母表示P < 0.05水平差异显著。The data in the table are mean ± standard error. Different lowercase letters after data in the same column indicate significant difference at P < 0.05 level.

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表3扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域冬小麦灌浆期旗叶光响应曲线参数
Table3.Light response curve parameters of flag leaves of winter wheat at filling stages in different intercropping areas under different canopy-shading degrees of almond-winter wheat intercropping system

处理 Treatment	间作小麦区域 Area of intercropped wheat	最大净光合速率 Pmax (μmol·m-2·s-1)	光饱和点 LSP (μmol·m-2·s-1)	暗呼吸速率 Rd (μmol·m-2·s-1)	光补偿点 LCP (μmol·m-2·s-1)	表观量子 效率 α
轻度遮阴 Light shading	近冠区?Near canopy	18.26±1.65a	1722.29±153.41a	-1.31±0.12c	15.60±1.46b	0.05b
	远冠区?Far from canopy	20.25±1.96a	1794.60±124.12a	-2.10±0.16a	31.26±2.18a	0.06ab
重度遮阴 Heavy shading	近冠区?Near canopy	10.46±0.77b	1366.30±113.14b	-1.73±0.14b	29.26±2.06a	0.07a
	远冠区?Far from canopy	11.44±1.01b	1474.88±126.89b	-1.73±0.15b	31.98±2.78a	0.06ab
单作小麦?Monocultured wheat (CK)		23.98±2.76a	1755.39±146.21a	-2.21±0.19a	37.51±2.14a	0.05b
表内数据为平均值±标准误, 同列数据后不同小写字母表示P < 0.05水平差异显著。Pmax: maximum net photosynthetic rate; LSP: light saturation point; Rd: dark respiration rate; LCP: light compensation point; α: initial quantum efficiency. The data in the table are mean ± standard error. Different lowercase letters after data in the same column indicate significant difference at P < 0.05 level.

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表4扁桃-冬小麦间作系统不同树冠截光程度下不同间作区域的冬小麦灌浆期旗叶荧光参数
Table4.Effects of canopy-shading degrees on fluorescence parameters of flag leaves of winter wheat in different intercropping areas of almond-winter wheat intercropping system

处理 Treatment	间作小麦区域 Area of intercropped wheat	PSⅡ最大光化学效率 Fv/Fm	PSⅡ实际光化学效率 ΦSPⅡ	光化学猝灭系数 qP	非光化学猝灭系数 NPQ
轻度遮阴 Light shading	近冠区?Near canopy	0.853±0.04a	0.299±0.02a	0.368±0.02b	0.499±0.02c
	远冠区?Far from canopy	0.836±0.05a	0.307±0.02a	0.384±0.01b	0.482±0.02c
重度遮阴 Sever shading	近冠区?Near canopy	0.848±0.05a	0.245±0.01b	0.295±0.01c	0.633±0.04a
	远冠区?Far from canopy	0.831±0.04a	0.248±0.01b	0.314±0.01c	0.527±0.03b
单作小麦?Monocultured wheat (CK)		0.857±0.03a	0.311±0.02a	0.509±0.03a	0.401±0.02d
表内数据为平均值±标准误, 同列数据后不同小写字母表示P < 0.05水平差异显著。Fv/Fm: maximal efficiency of PSⅡ photochemistry; ΦSPⅡ: effective quantum yield of PSⅡ photochemistry; qP: photochemical quenching; NPQ: non-photochemical quenching. The data in the table are mean ± standard error. Different lowercase letters after data in the same column indicate significant difference at P < 0.05 level.

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表5不同树冠截光程度对扁桃-冬小麦间作系统不同间作区域小麦产量及组成成分的影响
Table5.Effect of shading degree on yield and its' components of winter wheat in different intercropping area of almond-winter wheat intercropping

处理 Treatment	间作小麦区域 Area of intercropped wheat	产量 Yield (t·hm-2)	千粒重 1000-grain weight (g)	穗粒数 Grains number per spike	有效穗数 Effective panicle number (×104 panicles·hm-2)
轻度遮阴 Light shading	近冠区?Near canopy	5.05±0.41b	37.73±2.67b	25.78±1.43a	610.5±45.21ab
	远冠区?Far from canopy	5.63±0.34a	43.04±2.35a	24.27±1.67a	609.0±34.27ab
重度遮阴 Sever shading	近冠区?Near canopy	1.98±0.13c	33.46±3.01c	18.44±1.24b	368.5±32.12c
	远冠区?Far from canopy	2.26±0.15c	33.45±3.12c	14.96±1.11c	509.0±35.67b
单作小麦?Monocultured wheat (CK)		5.91±0.24a	39.01±2.97b	25.87±2.45a	663.21±52.34a
表内数据为平均值±标准误, 同列数据后不同小写字母表示P < 0.05水平差异显著。The data in the table are mean ± standard error. Different lowercase letters after the data in the same column indicate significant difference at P < 0.05.

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