席念勋3,
张大勇2
1.黄冈师范学院生物与农业资源学院/经济林木种质改良与资源综合利用湖北省重点实验室 黄冈 438000
2.北京师范大学生命科学学院/生物多样性与生态工程教育部重点实验室 北京 100875
3.中山大学生命科学学院/有害生物控制与资源利用国家重点实验室 广州 510275
基金项目: 国家自然科学基金项目31421063
国家自然科学基金项目41671040
高等学校学科创新引智计划B13008
黄冈师范学院培育基金项目201615703
详细信息
作者简介:朱莉, 主要研究方向为农业生态学及微生物生态学。E-mail:zhuli@hgnu.edu.cn
中图分类号:Q149计量
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出版历程
收稿日期:2019-12-10
录用日期:2020-04-13
刊出日期:2020-07-01
Effect of root competition on growth and resource allocation of two spring wheat varieties cultivated at different times
ZHU Li1, 2,,,XI Nianxun3,
ZHANG Dayong2
1. College of Biology and Agricultural Resources, Huanggang Normal University/Hubei Key Laboratory of Economic Forest Germplasm Improvement and Resources Comprehensive Utilization, Huanggang 438000, China
2. College of Life Sciences, Beijing Normal University/Key Laboratory for Biodiversity Science and Ecological Engineering, Ministry of Education, Beijing 100875, China
3. College of Life Sciences, Sun Yat-Sen University/State Key Laboratory of Biocontrol and Resource Utilization, Guangzhou 510275, China
Funds: the National Natural Science Foundation of China31421063
the National Natural Science Foundation of China41671040
the Program of Introducing Talents of Discipline to Universities of ChinaB13008
Huanggang Normal University High-level Cultivation Project201615703
More Information
Corresponding author:ZHU Li, E-mail:zhuli@hgnu.edu.cn
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摘要
摘要:植物相同基因型个体之间竞争地下资源会导致根系增生，这被称为根系竞争的“公地悲剧”现象。减少或者消除作物的根系增生有望提高种子产量。但人工育种是否已经无意识地影响了作物的根系竞争效应，目前研究尚不足。本研究选取春小麦古老品种‘和尚头’和现代品种‘92-46’开展温室盆栽试验，通过设置尼龙隔离、无竞争塑料隔离和有竞争塑料隔离，并设置1倍营养和2倍营养两个营养水平，分析根系竞争对春小麦各器官生物量的影响，并采用标准化主轴回归（SMA）方法分析根系竞争对资源分配的影响。结果表明，与无竞争塑料隔离相比，尼龙隔离的根系生物量无显著变化，但茎叶生物量和总生物量显著提高。尼龙隔离的古老品种‘和尚头’种子生物量与无竞争塑料隔离无显著差异，而现代品种‘92-46’的种子生物量显著高于无竞争塑料隔离。2倍营养-有竞争塑料隔离与1倍营养-无竞争塑料隔离相比，根系生物量和种子生物量无显著差异，但茎叶生物量和总生物量显著降低。标准化主轴回归（SMA）分析发现，根系竞争对‘和尚头’的根系资源分配无显著影响，根系竞争显著降低了‘92-46’的根系资源分配但降低程度较小（尼龙隔离相比无竞争塑料隔离）；根系竞争对两个品种的茎叶资源分配和种子资源分配均无显著影响。另外，本研究还发现：‘和尚头’的根系生物量和茎叶生物量大于‘92-46’，而种子生物量小于‘92-46’。除1倍营养-无竞争塑料隔离处理，其他隔离处理下‘和尚头’的根系资源分配与‘92-46’无显著差异；且‘和尚头’的茎叶资源分配大于‘92-46’，种子资源分配小于‘92-46’。总之，现代品种‘92-46’可能发展出一定程度的合作行为，通过根系分泌物介导识别邻株根系并减少根系资源分配；现代品种‘92-46’资源更多分配到繁殖生长，而古老品种‘和尚头’更多分配到茎叶生长。
关键词:达尔文农学/
“公地悲剧”/
根系竞争/
合作行为/
春小麦品种/
生物量/
资源分配
Abstract:Belowground competition among individuals of the same genotype can result in root proliferation, a pattern called "tragedy of the commons". Reducing or eliminating root proliferation could increase seed yield. However, it is not clear whether root competition effect has been influenced by breeding. A greenhouse pot experiment was conducted using an old spring wheat landrace 'Monkhead' and a modern cultivar '92-46'. Root interactions were set up by adding a mesh partition or a plastic partition between two plants from the same variety. Root competition was also encouraged by placing two plants on the same side of the plastic partition. There were two nutrition treatments: 1 time nutrition application and 2 times nutrition application. We examined the effect of root competition by comparing biomass and resource allocation patterns among the partition treatments. Biomass data showed that plants in the 'mesh partition' treatment produced similar root biomass to those in the 'plastic partition' treatment without competition, but their shoot biomass and total biomass were higher. 'Monkhead' plants in the 'mesh partition' treatment produced similar seed biomass to those in the 'plastic partition' treatment without competition, whereas seed biomass of '92-46' plants in the 'mesh partition' treatment was higher. Root biomass and seed biomass in the '2 times nutrition-plastic partition with competition' treatment were similar to those in the '1 time nutrition-plastic partition without competition' treatment, but their shoot biomass and total biomass were lower. A standardized major axis regression analysis (SMA) showed that root allocation of 'Monkhead' was not affected by root competition, but root allocation of '92-46' was decreased slightly ('mesh partition' vs. 'plastic partition without competition'). Root competition had no effect on shoot allocation and seed allocation for both varieties. In addition, Our results revealed composition differences between two varieties. 'Monkhead' produced more roots and shoots, but less seeds than '92-46'. 'Monkhead' also allocated more resources to shoots and less to seeds than '92-46', whereas root allocation was similar between two varieties in most partition sets except for '1 time nutrition-plastic partition without competition'. We conclude that the presence of roots from a neighboring plant does not result in 'the tragedy of the commons' pattern in either 'Monkhead' or '92-46', but affect the resource allocation pattern in '92-46' to some extent. Modern '92-46' is somehow able to develop restraining and cooperative behaviour characteristics in the presence of roots from neighboring plants, whereas 'Monkhead' does not show any responses. The differences in resource allocation patterns between two varieties support Darwinian agriculture, which suggests that in order to enhance yield, crop breeding has consciously selected varieties associated with lower competitiveness.
Key words:Darwinian agriculture/
Tragedy of the commons/
Root competition/
Cooperation/
Spring wheat varieties/
Biomass/
Resource allocation
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图1不同隔离处理示意图
Figure1.Illustration for different partition treatments
下载: 全尺寸图片幻灯片
图2尼龙隔离和无竞争塑料隔离下春小麦品种‘和尚头’和‘92-46’的生物量
柱状图显示均值±标准误差。*表示尼龙隔离和无竞争塑料隔离在0.05水平(双侧)上差异显著。
Figure2.Biomasses of spring wheat varieties 'Monkhead' and '92-46' under mesh partition and plastic partition without root competition
The data are mean ± SE. * means a significant difference between mesh partition and plastic partition without competition at 0.05 level (bilateral).
下载: 全尺寸图片幻灯片
图3根系竞争处理和无根系竞争处理下春小麦品种‘和尚头’和‘92-46’的生物量
柱状图显示均值±标准误差。*表示根系竞争处理和无根系竞争处理在0.05水平(双侧)上差异显著。
Figure3.Biomasses of spring wheat varieties 'Monkhead' and '92-46'with and without root competition
The data are mea ± SE. * means a significant difference between with and without root competition treatments at 0.05 level (bilateral).
下载: 全尺寸图片幻灯片表1根系竞争处理和无根系竞争处理下春小麦生长条件比较
Table1.Comparison of growing conditions of wheat plants with root competition and without root competition treatments
| 处理 Treatment | 单株可获得营养量 Available nutrient per plant (mg) | 单株可利用生长空间 Available space per plant | 每盆种植密度 Planting density per pot | 根系竞争 Root competition |
| 根系竞争处理(2倍营养-有竞争塑料隔离) Root competition (2 times nutrition-plastic partition with root competition) | 5.5 (4.25*) | 1/2花盆体积 1/2 pot volume | 2株 2 plants | 是 Yes |
| 无根系竞争处理(1倍营养-无竞争塑料隔离) No root competition (1 time nutrition-plastic partition without root competition) | 5.5 | 1/2花盆体积 1/2 pot volume | 2株 2 plants | 否 No |
| *表示该处理单株实际获得营养量, 是根据该处理实际所施加水溶液的浓度、体积、相应空间内小麦株数计算所得。* means actual amount of nutrition obtained by this individual plant, which was calculated according to the actual concentration and volume applied in this treatment and individual number in the corresponding space. | ||||
下载: 导出CSV表2尼龙隔离和无竞争塑料隔离下春小麦品种‘和尚头’和‘92-46’各器官生物量与全株总生物量之间的标准主轴回归分析
Table2.Standardized major axis regressions of biomass between different organs and whole plant for spring wheat varieties 'Monkhead' and '92-46' under mesh partition and plastic partition without competition
| 回归 Regression | 处理Treatment | 斜率 Slope (95% CI) | 截距 Intercept (95% CI) | R2 | P | |
| 品种Variety | 隔离Partition | |||||
| 根系-全株 Root-whole plant | 和尚头Monkhead | MP | 0.967a (0.748, 1.250) | –0.763ab (–0.947, –0.579) | 0.725 | < 0.001 |
| PP | 1.394a (1.138, 1.708) | –1.017ab (–1.199, –0.835) | 0.777 | < 0.001 | ||
| 92-46 | MP | 1.215a (1.028, 1.435) | –1.191b (–1.316, –1.066) | 0.886 | < 0.001 | |
| PP | 1.034a (0.904, 1.183) | –0.951a (–1.024, –0.877) | 0.926 | < 0.001 | ||
| 种子-全株 Seed-whole plant | 和尚头Monkhead | MP | 4.187a (2.825, 6.208) | –3.579b (–4.822, –2.336) | 0.335 | 0.008 |
| PP | 3.018a (1.902, 4.788) | –2.426b (–3.353, –1.498) | 0.126 | 0.135 | ||
| 92-46 | MP | 0.954b (0.787, 1.158) | –0.579a (–0.693, –0.465) | 0.846 | < 0.001 | |
| PP | 1.162b (0.948, 1.424) | –0.712a (–0.838, –0.586) | 0.828 | < 0.001 | ||
| 茎叶-全株 Shoot-whole plant | 和尚头Monkhead | MP | 0.939a (0.820, 1.076) | –0.081a (–0.175, 0.013) | 0.924 | < 0.001 |
| PP | 1.136a (0.921, 1.401) | –0.219ab (–0.372, –0.066) | 0.830 | < 0.001 | ||
| 92-46 | MP | 1.026a (0.969, 1.087) | –0.193b (–0.230, –0.157) | 0.986 | < 0.001 | |
| PP | 1.040a (0.937, 1.154) | –0.213b (–0.271, –0.156) | 0.955 | < 0.001 | ||
| MP:尼龙隔离; PP:无竞争塑料隔离。不同字母表示处理之间的斜率或者截距在0.05水平上差异显著。MP: mesh partition; PP: plastic partition without competition. Slopes and intercepts with different letters are significantly different among treatments at 0.05 levels. | ||||||
下载: 导出CSV表3根系竞争处理和无根系竞争处理下‘和尚头’和‘92-46’各器官生物量与全株总生物量之间的标准主轴回归分析
Table3.Standardized major axis regressions of biomass between different organs and whole plant for spring wheat varieties 'Monkhead' and '92-46' with and without root competition
| 回归 Regression | 处理Treatment | 斜率 Slope (95% CI) | 截距 Intercept (95% CI) | R2 | P | |
| 品种Variety | 根系竞争 Root competition | |||||
| 根系-全株 Root-whole plant | 和尚头 Monkhead | RC | 1.522ab (0.856, 2.705) | –1.016a (–1.508, –0.523) | 0.445 | 0.035 |
| NRC | 1.696a (1.375, 2.092) | –1.227a (–1.458, –0.997) | 0.933 | < 0.001 | ||
| 92-46 | RC | 1.184ab (0.777, 1.804) | –0.985a (–1.170, –0.800) | 0.717 | 0.002 | |
| NRC | 0.998b (0.776, 1.283) | –0.957a (–1.077, –0.838) | 0.903 | < 0.001 | ||
| 种子-全株 Seed-whole plant | 和尚头 Monkhead | RC | 4.706a (2.703, 8.190) | –3.226b (–4.687, –1.765) | 0.489 | 0.025 |
| NRC | 4.081a (2.137, 7.794) | –3.224b (–5.046, –1.402) | 0.278 | 0.118 | ||
| 92-46 | RC | 1.085b (0.854, 1.378) | –0.614a (–0.708, –0.520) | 0.912 | < 0.001 | |
| NRC | 1.219b (0.916, 1.622) | –0.764a (–0.931, –0.597) | 0.874 | < 0.001 | ||
| 茎叶-全株 Shoot-whole plant | 和尚头 Monkhead | RC | 0.841a (0.706, 1.003) | –0.055a (–0.134, 0.024) | 0.953 | < 0.001 |
| NRC | 0.861a (0.679, 1.092) | –0.029a (–0.162, 0.104) | 0.913 | < 0.001 | ||
| 92-46 | RC | 1.013a (0.807, 1.272) | –0.217b (–0.300, –0.133) | 0.921 | < 0.001 | |
| NRC | 0.993a (0.911, 1.084) | –0.172b (–0.213, –0.131) | 0.989 | < 0.001 | ||
| RC:根系竞争处理; NRC:无根系竞争处理。不同字母表示处理之间的斜率或者截距在0.05水平上差异显著。RC: root competition; NRC: no root competition. Slopes and intercepts with different letters are significantly different among treatments at 0.05 levels. | ||||||
下载: 导出CSV参考文献
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