Effects of intercropping oat with different crops on the community structure of soil nematodes and crop yields
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摘要: 间作是土地可持续利用的重要种植模式。土壤线虫作为土壤健康的指示生物, 可揭示地下生态系统的食物网功能。为探究燕麦不同间作模式对产量和土壤线虫群落的影响, 本试验分别设置燕麦单作、燕麦||大豆、燕麦||赤小豆、燕麦||马铃薯和燕麦||甘薯5个处理, 分析燕麦不同间作模式对作物产量以及土壤线虫的数量、多样性和群落结构的影响, 以筛选出较优的燕麦间作模式。结果表明: 相比单作, 间作模式在土地利用率和作物总产量方面有一定的优势, 其中燕麦||马铃薯间作模式最佳, 土地当量比(1.36)最高, 其次为燕麦||大豆间作模式, 土地当量比为1.29。4种燕麦间作模式均显著(P<0.05)降低了植物寄生线虫的相对丰度, 提高了食微线虫相对丰度, 优化了土壤线虫群落结构。燕麦||大豆间作模式下食微线虫比例最高(88.42%), 植物寄生线虫比例最低(6.31%), 且在多项生态指数中(瓦斯乐斯卡指数、线虫通路指数、多样性指数和均匀度指数)显著高于燕麦单作(P<0.05)。而燕麦||赤小豆间作模式下富集指数最高, 植食性线虫成熟度指数最低, 土壤线虫群落受干扰程度最低。速效钾与线虫通路指数呈极显著正相关(r=0.722**), 速效钾含量的提升可以使土壤有机质分解主要依靠细菌分解途径。综上, 间作种植模式可以优化土壤线虫群落结构, 改善土壤生态环境, 提高产量; 燕麦||大豆是该试验条件下最佳的间作模式。Abstract: Intercropping is an important planting method for the sustainable use of land resources, and soil nematodes are an indicator of soil health, which can reveal the function of food webs in underground ecosystems. To explore the effects of different intercropping patterns of oats on yield and soil nematode communities, five treatments were set up: oat monoculture, and intercropping systems of oat||soybean, oat||rice bean, oat||potato, and oat||sweet potato. The effects of different intercropping patterns of oats on crop yield and the number, diversity, and community structure of soil nematodes were analyzed to determine the best intercropping pattern. The results showed that, compared with monoculture, intercropping had certain advantages in terms of land utilization rate and total crop yield. The intercropping of oats and potatoes had the highest land equivalent ratio (1.36), Followed by the intercropping of oats and soybeans, the land equivalent ratio is 1.29. A total of 39 genera were identified, including 12 bacterivorous nematodes, 4 fungivorous nematodes, 13 plant-parasitic nematodes, and 10 omnivorous predatory nematodes. The four oat intercropping patterns significantly reduced the relative abundance of plant-parasitic nematodes, increased the relative abundance of beneficial nematodes (bacterivorous and fungivorous), and optimized the soil nematode community structure. Among them, the best intercropping pattern was oat and soybean, with the highest proportion of microbial-feeding nematodes (88.42%) and the lowest proportion of plant-parasite nematodes (6.31%). Additionally, the intercropping pattern of oat and soybean was significantly higher than that of oat monoculture (P<0.05) in multiple ecological indexes (Wasilewska index, nematode channel ratio, Shannon-Wiener index, and enrichment index), and was also the best among the four intercropping patterns. The enrichment index of oat and rice bean intercropping was the highest, and the plant parasite index was the lowest, indicating that the soil nematode community was the least disturbed. The enrichment index and structure index of each cropping pattern were lower than 50, and the soil nematode food web was in quadrant D, indicating that the soil environment was in a stressed state, the food web was degraded, and the soil nematode community was unstable at the experiment location. Pearson correlation analysis showed that soil nematode community ecological indices were correlated with soil physical and chemical properties. Organic matter was significantly negatively correlated with the evenness index (r=−0.635, P<0.05), significantly negatively correlated with the Shannon-Wiener index (r=−0.641, P<0.01), and significantly positively correlated with the plant parasitic index (r=0.633, P<0.05). There was a significant positive correlation between available K content and the nematode channel ratio (r=0.722, P<0.01), increasing available K content could change the decomposition pathway of soil organic matter and contribute more to the decomposition of soil organic matter. In conclusion, intercropping can optimize the soil nematode community structure, improve the soil ecological environment, and increase crop yield. Among the four intercropping patterns, oat and soybean intercropping were the best method, not only having the highest crop yield but also the most beneficial nematode community structure for soil health, which showed the strong production and ecological advantages of traditional Gramineae and Leguminosae intercropping.
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图 1 燕麦不同间作模式对土壤线虫总数和各营养类群相对丰度的影响
O: 燕麦单作; O||S: 燕麦||大豆; O||RB: 燕麦||赤小豆; O||P: 燕麦||马铃薯; O||SP: 燕麦||甘薯。不同小写字母表示不同种植模式间差异显著(P<0.05)。O: oat monoculture; O||S: oat||soybean; O||RB: oat||rice beans; O||P: oat||potato; O||SP: oat||sweet potato. Different lowercase letters indicate significant differences among different cropping patterns (P<0.05).
Figure 1. Effects of different intercropping patterns of oat on total number and relative abundance of nutrient groups of soil nematodes
图 2 燕麦不同间作模式的土壤线虫食物网结构富集图
O: 燕麦单作; O||S: 燕麦||大豆; O||RB: 燕麦||赤小豆; O||P: 燕麦||马铃薯; O||SP: 燕麦||甘薯。不同小写字母表示不同种植模式间差异显著(P<0.05)。O: oat monoculture; O||S: oat||soybean; O||RB: oat||rice beans; O||P: oat||potato; O||SP: oat||sweet potato. Different lowercase letters indicate significant differences among different cropping patterns (P<0.05).
Figure 2. Enrichment diagram of food web structure of soil nematodes in different intercropping patterns of oat
图 3 土壤线虫生态指数与土壤理化性质相关性热图
*: 显著相关(P<0.05 ); **: 极显著相关(P<0.01 )。Phosphatase: 磷酸酶; Sucrase: 蔗糖酶; Urease: 脲酶; Organic matter: 有机质; Available K: 速效钾; Available P: 有效磷; WI: 瓦斯乐斯卡指数; NCR: 线虫通路指数; RS: 丰富度指数; H′: 多样性指数; J: 均匀度指数; MI: 自由生活线虫成熟度指数; PPI: 植食性线虫成熟度指数; EI: 富集指数; SI: 结构指数。*: significant correlation at P<0.05; **: significant correlation at P<0.01. WI: Wasilewska index; NCR: nematode channel ratio; RS: species richness index; H′: Shannon-Wiener index; J: evenness index; MI: maturity index; PPI: plant parasite index; EI: enrichment index; SI: structure index.
Figure 3. Heatmap of correlation between soil nematode ecological index and soil physicochemical properties
表 1 燕麦不同间作模式对作物经济产量和土地当量比的影响
Table 1. Effects of different intercropping patterns of oat on crop economic yields and land equivalent ratios
种植模式
Cropping pattern产量 Yield (kg∙hm−2) 土地当量比
Land equivalent
ratio (LER)燕麦 Oat 大豆 Soybean 赤小豆 Rice bean 马铃薯 Potato 甘薯 Sweet potato 单作 Monoculture 2560.15±259.24a 3175.79±246.23a 2797.40±36.58a 26 245.60±2263.32a 25 530.70±997.06a 燕麦||大豆 Oat||soybean 1399.34±109.59c 2351.76±62.40b 1.29 燕麦||赤小豆 Oat||rice bean 1233.96±81.45c 1923.98±107.11b 1.17 燕麦||马铃薯 Oat||potato 1710.14±117.41b 18 087.20±2371.49b 1.36 燕麦||甘薯 Oat||sweet potato 1276.31±75.41c 16 572.20±2628.86b 1.15 间作模式中的作物产量是作物所占间作面积的产量, 同列不同小写字母表示种植模式间差异显著(P<0.05)。The crop yield in the intercropping mode is the yield of the intercropping area occupied by the crop. Different lowercase letters in the same column indicate significant differences among cropping patterns (P<0.05). 
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表 2 燕麦不同间作模式土壤的线虫群落组成及其相对丰度
Table 2. Soil nematode community composition and relative abundances in different intercropping patterns of oat
营养类群
Trophic
group科
Family属
Genusc-p值
Colonizer-persister
(c-p) value燕麦单作
Oat
monoculture燕麦||大豆
Oat||soybean燕麦||赤小豆
Oat||rice bean燕麦||马铃薯
Oat||potato燕麦||甘薯
Oat||sweet
potato食细菌线虫
Bacterivores小杆科 Rhabditidae 小杆属 Rhabditis 1 ++ ++ ++ ++ ++ 头叶科 Cephalobidae 头叶属 Cephalobus 2 +++ +++ ++ +++ +++ 真头叶属 Eucephalobus 2 ++ ++ ++ ++ ++ 丽突属 Acrobeles 2 ++ ++ ++ ++ + 拟丽突属 Acrobeloides 2 ++ 鹿角唇叶属 Cervidellus 2 ++ ++ ++ ++ ++ 绕线科 Plectidae 绕线属 Plectus 2 ++ ++ ++ ++ ++ 拟绕线属 Anaplectus 2 + ++ ++ ++ 威尔斯属 Wilsonema 2 ++ ++ ++ 畸头叶科 Teratocephalidae
畸头属 Teratocephalus 3 ++ ++ ++ ++ 棱咽科 Prismatolaimidae
棱咽属 Prismatolaimus 3 ++ ++ ++ ++ ++ 无咽科 Alaimidae
无咽属 Alaimus 4 ++ ++ ++ ++ ++ 食真菌线虫
Fungivores滑刃科
Aphelenchoididae滑刃属 Aphelenchoides 2 ++ ++ ++ ++ ++ 真滑刃科 Aphelenchidae
真滑刃属 Aphelenchus 2 +++ +++ +++ +++ ++ 膜皮科 Diphtherophoridae
膜皮属 Diphtherophora 3 ++ ++ ++ ++ ++ 细齿科 Leptonchidae
垫咽属 Tylencholaimus 4 ++ ++ ++ ++ ++ 植物寄生线虫
Plant-parasites垫刃科 Tylenchidae
垫刃属 Tylenchus 2 + 剑尾垫刃属 Malenchus 2 ++ ++ ++ + + 丝尾垫刃属 Filenchus 2 + 野外垫刃属 Aglenchus 2 + + 针科 Paratylenchidae
针属 Paratylenchus 2 ++ ++ + + 锥垫刃科 Tylodoridae
头垫刃属 Cephalenchus 3 ++ ++ + 锥科 Dolichodoridae
矮化属 Tylenchorhynchus 3 + 鞘科 Hemicycliophoridae
鞘属 Hemicycliophora 3 + ++ + 短体科 Pratylenchidae
短体属 Pratylenchus 3 ++ ++ ++ ++ 纽带科 Hoplolaimidae
螺旋属 Helicotylenchus 3 ++ + 毛刺科 Trichodoridae
毛刺属 Trichodorus 4 + ++ ++ ++ + 长针科 Longidoridae
长针属 Longidorus 5 ++ ++ ++ 剑属 Xiphinema 5 + ++ ++ ++ 杂食线虫
Omnivores-predators倒齿科 Anatonchidae
等齿属 Miconchus 4 ++ ++ ++ + 三孔科 Tripylidae
三孔属 Tripyla 4 + ++ ++ + 单齿科 Monochidae
锯齿属 Prionchulus 4 + ++ ++ ++ ++ 锉齿属 Mylonchulus 4 + + 矛线科 Dorylaimidae 前矛线属 Prodorylaimus 4 + ++ + 真矛线属 Eudorylaimus 4 ++ ++ ++ ++ 中矛线属 Mesodorylaimus 4 ++ + 螯属 Pungentus 4 + ++ + 鄂针科 Belondiridae 鄂针属 Belondira 5 ++ ++ ++ + + 缢咽属 Axonchium 5 ++ ++ ++ +++表示丰度>10%, 为优势属; ++表示丰度为1%~10%, 为常见属; +表示丰度<1%, 为稀有属。+++ indicates abundance >10%, the dominant genus; ++ indicates that the abundance is between 1%−10%, the common genus; + indicates abundance <1%, the rare genus.
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表 3 燕麦不同间作模式对土壤线虫群落生态指数的影响
Table 3. Effects of different intercropping patterns of oat on ecological indices of soil nematode communities
生态指数
Ecological indice燕麦单作
Oat monoculture燕麦||大豆
Oat||soybean燕麦||赤小豆
Oat||rice bean燕麦||马铃薯
Oat||potato燕麦||甘薯
Oat||sweet potato瓦斯乐斯卡指数 Wasilewska index 5.78±0.21b 14.68±3.51a 12.17±2.25a 10.15±2.19ab 14.28±4.33a 线虫通路指数 Nematode channel ratio 0.55±0.03b 0.71±0.01a 0.55±0.01b 0.71±0.02a 0.69±0.08a 丰富度指数 Species richness index 4.68±0.39a 3.11±0.18c 3.93±0.28b 3.58±0.06b 3.94±0.57b 多样性指数 Shannon-Wiener index 4.94±0.21d 6.27±0.10a 5.72±0.17bc 5.26±0.43cd 5.89±0.30ab 均匀度指数 Evenness index 0.84±0.03b 1.07±0.21a 1.02±0.04a 0.90±0.09b 0.99±0.10a 自由生活线虫成熟度指数 Maturity index 33.54±23.55a 21.55±1.91a 13.39±1.76a 17.41±0.82a 20.05±1.30a 植食性线虫成熟度指数 Plant parasite index 4.54±0.96a 2.42±0.75ab 1.36±0.34b 4.02±2.28a 2.08±1.18ab 同行不同小写字母表示不同处理间差异显著(P<0.05)。Different lowercase letters in the same row indicate significant differences among different cropping patterns atP<0.05 level.
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