关键词:棉花; 嫁接; 叶片; 衰老; 根-冠互作 Effect of Root-shoot Interaction on Cotton Leaf Senescence ZHANG Qiao-Yu, WANG Yi-Ru, AN Jing, WANG Bao-Min, TIAN Xiao-Li* Center of Crop Chemical Regulation, College of Agricultural and Biotechnology, China Agricultural University / State Key Laboratory of Plant Physiology and Biochemistry, Beijing 100193, China Fund:This study was supported by the National Natural Science Foundation of China (31271629) AbstractResearches and practices associated with grafting indicate that the diversity of root-shoot interaction is universal in different physiological processes of plant. Our previous grafting study with CCRI41, an early senescence cotton cultivar, and SCRC22, a late senescence cultivar as materials found that shoot played a major role in mediating leaf senescence of cotton. In the present study, CCRI41 and another late senescence cotton cultivar CCRI49 were used to do standard grafting (I-grafting, one scion grafted onto one rootstock) and Y-grafting (two scions grafted onto one rootstock). Leaf senescence of grafts was induced by low potassium (K; 0.03 mmol L-1). Contrary to the grafting combination of CCRI41 and SCRC22, the combination of CCRI41 and CCRI49 showed that the role of root was more important than that of shoot for leaf senescence, as characterized by SPAD reading. Accordingly, the concentration of ZR+Z or iPA+iP in roots and the youngest fully expanded leaf and their delivery rate in xylem sap regarding I-grafting combinations with CCRI41 as rootstock were significantly lower than those with CCRI49 as rootstock in most of situations. However, the results of free ABA level were exactly opposite. With respect to the Y-grafting of CCRI41 and CCRI49, the results were similar to those of I-grafting, whereas the role of root was not absolutely predominant any more. The mechanism for different root-shoot interactions in cotton leaf senescence was discussed from the perspective of root-shoot-root distance communication.
图1 高钾条件下(2.5 mmol L-1) I型(A)和Y型(B)嫁接植株的倒四叶Fig. 1 The youngest fully expanded leaf of I grafts (A) and Y grafts (B) under sufficient potassium (2.5 mmol L-1)
图2 低钾条件下(0.03 mmol L-1) I型(A)和Y型(B)嫁接植株的倒四叶Fig. 2 The youngest fully expanded leaf of I grafts (A) and Y grafts (B) under potassium deficiency (0.03 mmol L-1)
表1 Table 1 表1(Table 1)
表1 供钾水平对I型和Y型嫁接各处理倒四叶SPAD值的影响 Table 1 Effect of potassium (K) supply on SPAD value of the youngest fully expanded leaf of cotton standard grafts (scion/rootstock) and Y grafts (scion+scion/rootstock)
嫁接方式 Grafting method
供K水平 K supply (mmol L-1)
嫁接处理 Combination of scion and rootstock
I型 Scion/rootstock
41/41
49/41
41/49
49/49
2.50
37.9 a
38.1 a
38.0 a
38.4 a
0.03
18.4 d
20.3 c
21.5 b
24.3 a
Y型scion+scion/rootstock
41+41/41
41+41/41
41+49/41
41+49/41
41+49/49
41+49/49
49+49/49
49+49/49
2.50
36.0 a
35.2 a
33.7 a
34.4 a
34.9 a
35.5 a
35.7 a
34.3 a
0.03
18.3 b
19.3 b
18.2 b
21.1 ab
20.4 ab
25.5 a
22.7 ab
23.5 ab
41: CCRI41, early senescence under K deficiency; 49: CCRI49, late senescence under K deficiency. The data within the same grafting method followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test. 41: 中棉所41; 49: 中棉所49。每列数据为粗体接穗的结果; 各嫁接方式不同嫁接处理平均值后的不同字母表示在0.05水平差异显著。
表1 供钾水平对I型和Y型嫁接各处理倒四叶SPAD值的影响 Table 1 Effect of potassium (K) supply on SPAD value of the youngest fully expanded leaf of cotton standard grafts (scion/rootstock) and Y grafts (scion+scion/rootstock)
表2 低钾条件下(0.03 mmol L-1)不同嫁接方式倒四叶SPAD值的砧木和接穗调控效应 Table 2 Effects of rootstock and scion on SPAD value of the youngest fully expanded leaf under low K+ (0.03 mmol L-1) condition (%)
嫁接方式 Grafting method
中棉所41 CCRI 41
中棉所49 CCRI 49
砧木效应 Rootstock effect
接穗效应 Sion effect
砧木效应 Rootstock effect
接穗效应 Sion effect
I型 One scion/one rootstock
-16.3
-11.6
16.5
10.3
Y型 Two scions/one rootstock
-8.5
-0.6
8.4
4.8
表2 低钾条件下(0.03 mmol L-1)不同嫁接方式倒四叶SPAD值的砧木和接穗调控效应 Table 2 Effects of rootstock and scion on SPAD value of the youngest fully expanded leaf under low K+ (0.03 mmol L-1) condition (%)
图3 供钾水平对I型嫁接各处理根系(R)和倒四叶(L)的ZR+Z浓度(ng g-1 FW)及嫁接位点上部(A)和下部(B)木质部汁液ZR+Z流量(ng plant-1 h-1)的影响 A: 充足供钾(2.5 mmol L-1); B: 低钾胁迫(0.03 mmol L-1)。同一供钾水平, 同一部位(指根系、叶片和木质部汁液)数据后的不同小写字母表示在0.05水平差异显著(n = 4)。Fig. 3 Effect of K deficiency on ZR+Z level of cotton standard grafts (scion/rootstock) A: sufficient K (2.5 mmol L-1). B: low K (0.03 mmol L-1). The ZR+Z concentration (ng g-1FW) in roots (R) and the youngest fully expanded leaf (L), and ZR+Z delivery rates (ng plant-124 h-1) in xylem above- (A) and below graft union (B) were determined. Means of the same sampling part (i.e. roots, leaf, and xylem sap collected both below and above the graft union) within the same K level followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test (n = 4).
图4 供钾水平对I型嫁接各处理根系(R)和倒四叶(L)的iPA+iP浓度(ng g-1 FW)及嫁接位点上(A)和下部(B)木质部汁液iPA+iP流量(ng plant-1 h-1)的影响 A: 充足供钾(2.5 mmol L-1); B: 低钾胁迫(0.03 mmol L-1)。同一供钾水平, 同一部位(指根系、叶片和木质部汁液)数据后的不同小写字母表示在0.05水平差异显著(n = 4)。Fig. 4 Effect of K deficiency on iPA+iP level of cotton standard grafts (scion/rootstock) A: sufficient K (2.5 mmol L-1). B: low K (0.03 mmol L-1). The iPA+iP concentration (ng g-1FW) in roots (R) and the youngest fully expanded leaf (L), and iPA+iP delivery rates (ng plant-124 h-1) in xylem above- (A) and below graft union (B) were determined. Means of the same sampling part (i.e. roots, leaf, and xylem sap collected both below and above the graft union) within the same K level followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test (n = 4).
图5 供钾水平对I型嫁接各处理根系(R)和倒四叶(L)的ABA浓度(ng g-1 FW)及嫁接位点上部(A)和下部(B)木质部汁液ABA流量(ng plant-1 h-1)的影响 A: 充足供钾(2.5 mmol L-1); B: 低钾胁迫(0.03 mmol L-1)。同一供钾水平, 同一部位(指根系、叶片和木质部汁液)数据后的不同小写字母表示在0.05水平差异显著(n = 4)。Fig. 5 Effect of K deficiency on ABA level of cotton standard grafts (scion/rootstock) A: sufficient K (2.5 mmol L-1). B: low K (0.03 mmol L-1). The ABA concentration (ng g-1 FW) in roots (R) and the youngest fully expanded leaf (L), and ABA delivery rates (ng plant-124 h-1) in xylem above- (A) and below graft union (B) were determined. Means of the same sampling part (i.e. roots, leaf, and xylem sap collected both below and above the graft union) within the same K level followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test (n = 4).
表3 低钾条件下(0.03 mmol L-1) I型嫁接倒四叶和接穗木质部汁液中激素水平的砧木和接穗效应 Table 3 Effect of rootstock and scion on phytohormone levels in the youngest fully expanded leaf and the xylem sap above graftunion for standard graftings (one scion/one rootstock) under low K+ (0.03 mmol L-1) condition (%)
植物激素 Phytohormone
中棉所41 CCRI 41
中棉所49 CCRI 49
砧木效应 Rootstock effect
接穗效应 Scion effect
砧木效应 Rootstock effect
接穗效应 Scion effect
倒四叶 The youngest fully expanded leaf
ZR+Z
-21.4
-9.2
25.9
9.1
iPA+iP
-10.9
-5.4
8.1
1.8
ABA
78.3
10.3
-47.7
-15.5
嫁接位点上部木质部汁液 The xylem sap above graft union
ZR+Z
-36.3
-17.3
50.0
15.6
iPA+iP
-36.0
-17.1
106.2
59.1
ABA
113.3
41.7
-39.1
-8.4
表3 低钾条件下(0.03 mmol L-1) I型嫁接倒四叶和接穗木质部汁液中激素水平的砧木和接穗效应 Table 3 Effect of rootstock and scion on phytohormone levels in the youngest fully expanded leaf and the xylem sap above graftunion for standard graftings (one scion/one rootstock) under low K+ (0.03 mmol L-1) condition (%)
图6 供钾水平对Y型嫁接处理根系(R)和倒四叶(L)的ZR+Z浓度(ng g-1 FW)及嫁接位点上部(A)和下部(B)木质部汁液ZR+Z流量 (ng plant-1 h-1)的影响 A:充足供钾(2.5 mmol L-1); B: 低钾胁迫(0.03 mmol L-1)。同一供钾水平, 同一部位(指根系、叶片和木质部汁液)数据后的不同小写字母表示在0.05水平差异显著(n = 4)。Fig. 6 Effect of K deficiency on ZR+Z level of cotton Y grafts (two scions/rootstock) A: sufficient K (2.5 mmol L-1); B: low K (0.03 mmol L-1). The ZR+Z concentration (ng g-1 FW) in roots (R) and the youngest fully expanded leaf (L), and ZR+Z delivery rates (ng plant-124 h-1) in xylem above- (A) and below graft union (B) were determined. Means of the same sampling part (i.e. roots, leaf, and xylem sap collected both below and above the graft union) within the same K level followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test (n = 4).
图7 供钾水平对Y型嫁接各处理根系(R)和倒四叶(L)的iPA+iP浓度(ng g-1 FW)及嫁接位点上部(A)和下部(B)木质部汁液iPA+iP流量(ng plant-1 h-1)的影响 A:充足供钾(2.5 mmol L-1); B: 低钾胁迫(0.03 mmol L-1)。同一供钾水平, 同一部位(指根系、叶片和木质部汁液)数据后的不同小写字母表示在0.05水平差异显著(n = 4)。Fig. 7 Effect of K deficiency on iPA+iP level of cotton Y grafts (two scions/rootstock) A: sufficient K (2.5 mmol L-1). B: low K (0.03 mmol L-1). The iPA+iP concentration (ng g-1 FW) in roots (R) and the youngest fully expanded leaf (L), and iPA+iP delivery rates (ng plant-124 h-1) in xylem above- (A) and below graft union (B) were determined. Means of the same sampling part (i.e. roots, leaf, and xylem sap collected both below and above the graft union) within the same K level followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test (n = 4).
图8 供钾水平对Y型嫁接各处理根系(R)和倒四叶(L)的ABA浓度(ng g-1 FW)及嫁接位点上部(A)和下部(B)木质部汁液ABA流量(ng plant-1 h-1)的影响 A: 充足供钾(2.5 mmol L-1); B: 低钾胁迫(0.03 mmol L-1)。同一供钾水平, 同一部位(指根系、叶片和木质部汁液)数据后的不同小写字母表示在0.05水平差异显著(n = 4)。Fig. 8 Effect of K deficiency on ABA level of cotton Y grafts (two scions/rootstock) A: sufficient K (2.5 mmol L-1); B: low K (0.03 mmol L-1). The ABA concentration (ng g-1 FW) in roots (R) and the youngest fully expanded leaf (L), and ABA delivery rates (ng plant-124 h-1) in xylem above- (A) and below graft union (B) were determined. Means of the same sampling part (i.e. roots, leaf, and xylem sap collected both below and above the graft union) within the same K level followed by the same letter are not significantly different at P < 0.05 according to Duncan’ s multiple range test (n = 4).
表4 低钾条件下(0.03 mmol L-1) Y型嫁接倒四叶和接穗木质部汁液中激素水平的砧木和接穗效应 Table 4 Effect of rootstock and scion on phytohormone levels in the youngest fully expanded leaf and the xylem sap above graf tunion for Y graftings (two scion/one rootstock) under low K+ (0.03 mmol L-1) condition (%)
植物激素 Phytohormone
中棉所41 CCRI 41
中棉所49 CCRI 49
砧木效应 Rootstock effect
接穗效应 Scion effect
砧木效应 Rootstock effect
接穗效应 Scion effect
倒4叶 The youngest fully expanded leaf
ZR+Z
-11.7
-1.9
72.5
45.8
iPA+iP
-9.0
-1.7
13.0
10.8
ABA
21.6
2.3
-24.1
-10.0
嫁接位点上部木质部汁液 The xylem sap above graft union
ZR+Z
-15.2
-8.7
21.3
20.7
iPA+iP
-26.3
-8.6
8.4
4.2
ABA
23.9
9.7
-18.1
-2.1
表4 低钾条件下(0.03 mmol L-1) Y型嫁接倒四叶和接穗木质部汁液中激素水平的砧木和接穗效应 Table 4 Effect of rootstock and scion on phytohormone levels in the youngest fully expanded leaf and the xylem sap above graf tunion for Y graftings (two scion/one rootstock) under low K+ (0.03 mmol L-1) condition (%)
图9 棉花叶片衰老中不同根-冠互作类型的根-冠-根通讯机制 41: 中棉所41; 49: 中棉所49; 22: 鲁棉研22; 41/22: 地上部为主型; 41/49: 根系为主型; 绿色上行箭头: 根-冠信号; 蓝色下行箭头: 冠-根信号; 箭头粗细代表信号强弱。Fig. 9 The diagram of root-shoot-root communication for different interactions between root and shoot in cotton leaf senescence induced by low potassium (0.03 mmol L-1) stress 41: CCRI41; 49: CCRI49; 22: SCRC22; 41/22: the role of shoot is more important; 41/49: the role of root is more important. Green up arrow: the signal of root-shoot. Blue down arrow: the signal of shoot-root. The thickness of arrow indicates signal strength.
4 结论以中41为砧木的处理, 其叶片中ZR+Z、iPA+iP的含量低于以中49为砧木的处理, ABA含量则相反, 表明根系对叶片衰老的作用较大。据分析, 中41和中49互相嫁接体(41/49和49/41)接穗叶片中的受体感知到来自砧木的缺钾信号后, 发出的冠-根信号对砧木输出CK和ABA的能力和/或木质部中CK和ABA的运输过程影响较小; 接穗叶片中CK和ABA的合成、代谢则可能发生类似于砧木品种的变化。 致谢: 感谢课题组王晔博士在试验过程中提供的帮助。 The authors have declared that no competing interests exist.
作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.
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, 王逸茹, 安静, 王保民, 田晓莉
表1 供钾水平对I型和Y型嫁接各处理倒四叶SPAD值的影响 Table 1 Effect of potassium (K) supply on SPAD value of the youngest fully expanded leaf of cotton standard grafts (scion/rootstock) and Y grafts (scion+scion/rootstock)
