Effects of intercropping with legumes on the activities of key enzymes of carbon and nitrogen metabolism in potato leaves and crop yield
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摘要:
马铃薯连作会产生诸多危害, 研究与豆类作物间作对马铃薯叶片碳氮代谢关键酶活性及作物产量的影响, 进而选择适宜的马铃薯栽培模式, 对提高农业生产效益、保护生态环境以及推动农业可持续发展有重要意义。本研究于2020年5月至2021年9月在宁夏海原县开展, 以马铃薯品种‘青薯9号’为材料, 采取随机区组试验, 设置马铃薯单作(IP)、大豆单作(IS)、蚕豆单作(IB)、马铃薯与大豆间作(PS)、马铃薯与蚕豆间作(PB) 5种种植模式, 测定马铃薯叶片碳代谢、氮代谢相关指标及作物产量。结果表明: 与豆类作物间作能显著提高马铃薯叶片碳氮代谢相关酶活性、增加群体产量, 提高土地利用率, 其中以马铃薯与大豆间作(PS)表现最佳。相关性分析表明, 碳氮代谢各指标均与产量呈极显著正相关, 碳代谢关键酶(蔗糖合成酶、蔗糖磷酸合成酶)与氮代谢关键酶(谷氨酰胺合成酶、硝酸还原酶)之间存在极显著正相关。主成分分析表明, 与豆类作物间作时, 可溶性糖、蔗糖磷酸合成酶、谷氨酰胺合成酶及硝酸还原酶对马铃薯产量的提升起主导作用。马铃薯与豆类间作土地当量比均大于1, 其中马铃薯与大豆间作(PS)两年平均土地当量比为1.20, 表现出显著的间作优势。综上, 马铃薯叶片碳氮代谢酶之间存在显著的正向影响, 且对产量形成有促进作用, 与豆类间作可显著提升马铃薯叶片碳氮代谢酶活性, 进而提高群体产量。在宁夏南部半干旱区, 为克服马铃薯连作障碍、促进农业可持续发展、提升经济效益, 可推广马铃薯与豆类间作模式, 以马铃薯与大豆间作最宜。
Abstract:As continuous cropping of potatoes produces many hazards, it is important to study the effects of intercropping with legumes on the activities of key enzymes of carbon and nitrogen metabolism in potato leaves and crop yields and to select suitable potato cultivation modes to improve the efficiency of agricultural production, protect the ecological environment, and promote the sustainable development of agriculture. We measured the activities of key enzymes relative to carbon and nitrogen metabolism of potato leaf, and crops yields. The results showed that intercropping with legumes could significantly increase the activities of carbon and nitrogen metabolism-related enzymes in potato leaves, increase the yield of the population, and improve the land utilization rate; intercropping between potato and soybean (PS) performed the best. Correlation analysis showed that all enzymes activities of carbon and nitrogen metabolism were positively correlated with yield and that there was a significant positive correlation between the key enzymes of carbon metabolism (sucrose synthase and sucrose phosphate synthase) and the key enzymes of nitrogen metabolism (glutamine synthase and nitrate reductase). Principal component analysis showed that soluble sugars, sucrose phosphate synthetase, glutamine synthetase, and nitrate reductase played dominant roles in improving potato yield during intercropping with legumes. The land equivalent ratios of intercropping between potatoes and legumes were all greater than 1, and the average two-year land equivalent ratio of intercropping between potato and soybean (PS) was 1.20, showing significant intercropping advantages. In conclusion, there is a significant positive correlation between potato leaf carbon and nitrogen metabolism enzymes, which promotes yield formation, and intercropping with beans can significantly enhance potato leaf carbon and nitrogen metabolism enzyme activity, thus improving group yield. In the semi-arid region of southern Ningxia, to overcome the obstacles of potato succession, promote sustainable agricultural development, and enhance economic benefits, we can promote intercropping modes of potatoes and beans, with intercropping of potato and soybean (PS) being the most suitable.
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Keywords:
- Potato /
- Legume /
- Intercropping /
- Carbon and nitrogen metabolism enzymes /
- Land equivalent ratio /
- Yield
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图 3 2020年和2021年与大豆和蚕豆间作对不同生育期马铃薯叶片氮代谢关键酶活性的影响
S: 苗期; B: 现蕾期; TF: 块茎形成期; TE: 块茎膨大期; M: 成熟期; IP: 马铃薯单作; PS: 马铃薯与大豆间作; PB: 马铃薯与蚕豆间作。同生育期不同小写字母表示不同处理间在P<0.05水平差异显著。S: seedling stage; B: budding stage; TF: tuber formation stage; TE: tuber expansion stage; M: maturity stage; IP: potato monoculture; PS: potato and soybean intercropping; PB: potato and faba bean intercropping. Different lowercase letters represent significant differences among treatments at P<0.05 level at the same growth stage.
Figure 3. Effect of intercropping with soybean and faba bean on activities of key enzymes of nitrogen metabolism in potato leaves at different growth stages in 2020 and 2021
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图 1 2020年和2021年试验区马铃薯生育期内气温和降雨量变化
Figure 1. Variations in temperature and precipitation during the potato growth periods in the experimental area in 2020 and 2021
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图 2 2020年和2021年与大豆和蚕豆间作对不同生育期马铃薯叶片碳代谢关键酶活性的影响
S: 苗期; B: 现蕾期; TF: 块茎形成期; TE: 块茎膨大期; M: 成熟期; IP: 马铃薯单作; PS: 马铃薯与大豆间作; PB: 马铃薯与蚕豆间作。同生育期不同小写字母表示不同处理间在P<0.05水平差异显著。S: seedling stage; B: budding stage; TF: tuber formation stage; TE: tuber expansion stage; M: maturity stage; IP: potato monoculture; PS: potato and soybean intercropping; PB: potato and faba bean intercropping. Different lowercase letters represent significant differences among treatments at P<0.05 level at the same growth stage.
Figure 2. Effect of intercropping with soybean and faba bean on activities of key enzymes of carbon metabolism in potato leaves at different growth stages in 2020 and 2021
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图 4 2020年和2021年与大豆和蚕豆间作对不同生育期马铃薯叶片可溶性糖和可溶性蛋白的影响
S: 苗期; B: 现蕾期; TF: 块茎形成期; TE: 块茎膨大期; M: 成熟期; IP: 马铃薯单作; PS: 马铃薯与大豆间作; PB: 马铃薯与蚕豆间作。同生育期不同小写字母表示不同处理间在P<0.05水平差异显著。S: seedling stage; B: budding stage; TF: tuber formation stage; TE: tuber expansion stage; M: maturity stage; IP: potato monoculture; PS: potato and soybean intercropping; PB: potato and faba bean intercropping. Different lowercase letters represent significant differences among treatments at P<0.05 level at the same growth stage.
Figure 4. Effect of intercropping with soybean and faba bean on contents of soluble sugar and soluble protein of potato leaves at different growth stages in 2020 and 2021
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图 5 马铃薯叶片碳氮代谢酶活性与产量的相关性分析
A: 可溶性糖; B: 可溶性蛋白; SS: 蔗糖合成酶; SPS: 蔗糖磷酸合成酶; GS: 谷氨酰胺合成酶; NR: 硝酸还原酶; Yield: 产量; *: P<0.05; **: P<0.01. A: soluble sugar; B: soluble protein; SS: sucrose synthase; SPS: sucrose phosphate synthase; GS: glutamine synthase; NR: nitrate reductase.
Figure 5. Correlation between key enzyme activities relative to carbon and nitrogen metabolism and potato yield
表 1 试验区土壤基础肥力
Table 1 Soil basic properties of the tested area
年份
YearpH 有机质
Organic matter
(g·kg−1)全氮
Total nitrogen
(g·kg−1)碱解氮
Available nitrogen
(mg·kg−1)速效磷
Available phosphorus
(mg·kg−1)速效钾
Available potassium
(mg·kg−1)2020 8.04 10.95 0.84 62.67 42.18 177.45 2021 8.07 10.42 1.05 58.42 41.29 165.17 
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表 2 2020年和2021年马铃薯与大豆和蚕豆间作对作物当量产量、土地当量比及种间相对竞争的影响
Table 2 Effects of intercropping of potato with soybean and faba bean on crop equivalent yield, land equivalent ratio, and relative interspaces competition in 2020 and 2021
年份
Year处理
Treatment当量产量
Equivalent yield (kg·hm−2)偏土地当量比
Partial land equivalent ratio土地当量比
Land equivalent ratio种间相对
竞争力
Relative interspecies competitiveness马铃薯
Potato大豆
Soybean蚕豆
Faba bean马铃薯
Potato大豆
Soybean蚕豆
Faba bean2020 SC 29 082.24±422.09a 4792.50±540.94a 3333.00±247.35a PS 16 570.78±105.80b 3039.19±105.79b 0.57 0.63 1.20 −0.13 PB 15 316.25±418.18c 2011.35±128.34b 0.53 0.60 1.13 −0.15 2021 SC 27 132.24±337.48a 4532.50±200.55a 3146.03±181.87a PS 15 615.78±100.60b 2816.69±134.53b 0.58 0.62 1.20 −0.09 PB 14 416.25±193.33c 1901.76±131.46b 0.53 0.60 1.14 −0.15 变异来源
Source of variationY ** NS NS T ** ** ** Y×T * NS NS SC: 单作(马铃薯、大豆和蚕豆单作分别为IP、IS和IB处理); PS: 马铃薯与大豆间作; PB: 马铃薯与蚕豆间作; Y: 年份; T: 处理; Y×T: 年份×处理。NS: 不显著; *: P<0.05水平差异显著; **: P<0.01水平差异显著。表格中同年同列不同小写字母表示处理间在P<0.05水平显著。SC: sole cropping (potato, soybean and faba bean monocropping are IP, IS and IB treatments, respectively); PS: potato and soybean intercropping; PB: potato and faba bean intercropping; Y: year; T: treatment; Y×T: year × treatment. NS: no significant difference; *: significant differences at P<0.05 level; **: significant differences at P<0.01 level. Different lowercase letters in the same column for the same year indicate significant differences among treatments at P<0.05 level.
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表 3 马铃薯叶片碳氮代谢关键酶活性与产量的主成分分析
Table 3 Principal component analysis of the activities of key enzymes of carbon and nitrogen metabolism in potato leaves in relation to yield
指标 Index PC1 PC2 PC3 可溶性糖 Soluble sugar 0.893 −0.255 −0.324 可溶性蛋白 Soluble protein 0.817 −0.494 0.266 蔗糖合成酶 Sucrose synthase 0.803 0.529 0.083 蔗糖磷酸合成酶 Sucrose phosphate synthase 0.935 0.176 −0.204 谷氨酰胺合成酶 Glutamine synthase 0.928 0.175 0.202 硝酸还原酶 Nitrate reductase 0.960 −0.217 −0.014 产量 Yield 0.974 0.092 0.022 特征值 Eigenvalue 0.895 0.706 0.266 方差贡献率 Variance contribution (%) 81.635 10.079 3.799 累计方差贡献率 Cumulative variance contribution (%) 81.635 91.714 95.513
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