不同水肥运筹对再生季稻根际土壤酶活性及微生物功能多样性的影响

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陈鸿飞, 庞晓敏, 张仁, 张志兴, 徐倩华, 方长旬, 李经勇, 林文雄. . 不同水肥运筹对再生季稻根际土壤酶活性及微生物功能多样性的影响. 作物学报, 2017, 43(10): 1507-1517
CHEN Hong-Fei, PANG Xiao-Min, ZHANG Ren, ZHANG Zhi-Xing, XU Qian-Hua, FANG Chang-Xun, LI Jing-Yong, LIN Wen-Xiong. . Effects of Different Irrigation and Fertilizer Application Regimes on Soil Enzyme Activities and Microbial Functional Diversity in Rhizosphere of Ratooning Rice. ACTA AGRONOMICA SINICA, 2017, 43(10): 1507-1517 复制到剪切板

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不同水肥运筹对再生季稻根际土壤酶活性及微生物功能多样性的影响

陈鸿飞^1,^*, 庞晓敏^1,^*, 张仁², 张志兴¹, 徐倩华³, 方长旬¹, 李经勇⁴, 林文雄^1,^*

¹福建农林大学生命科学学院 / 福建省农业生态过程与安全监控重点实验室, 福建福州350002

²福建省南平市建阳区农业局, 福建建阳353000

³福建省农业厅种植业技术推广总站, 福建福州 350001

⁴重庆市农业科学院, 重庆 402160

^* 通讯作者(Corresponding author): 林文雄, E-mail: lwx@fafu.edu.cn ^*同等贡献(Contributed equally to this work)

第一作者联系方式: 陈鸿飞, E-mail: hongfeichen2006@163.com

收稿日期:2017-03-06 接受日期:2017-05-10 网络出版日期:2017-06-09

基金:本研究由国家重点研发计划项目(2016YFD0300508), 福建省高校青年自然基金重点项目(JZ160435)和福建农林大学科技创新专项基金项目(CXZX2016076, CXZX2016077)资助

摘要

再生季水、肥管理措施是再生稻尤其是机收中低留桩再生稻获得高产稳产的一项关键措施, 明确再生季合理的水肥运筹对提高机收中低留桩再生稻低节位再生芽的萌发成穗具有重要意义。本文以杂交籼稻泸优明占为材料, 通过设置再生季分蘖期3个不同水肥处理盆栽试验, 即淹水灌溉不施用促苗肥(S处理)、干湿交替灌溉不施用促苗肥(G处理)和干湿交替灌溉施用促苗肥(GN处理), 研究不同水肥运筹对再生季稻腋芽、产量、根际土壤酶活性及微生物功能多样性的影响。结果表明, 再生季稻不同水肥处理对根际氧化还原电位和根际土壤酶有显著影响, 处理后10 d, 与S处理相比, G处理可以显著提高根际土壤的氧化还原电位和根际土壤多酚氧化酶、过氧化物酶、磷酸单酯酶、过氧化氢酶活性, 与G处理相比, GN处理可显著提高根际土壤多酚氧化酶、过氧化物酶、脲酶、蔗糖酶、磷酸单酯酶、过氧化氢酶活性。不同水肥调控下再生季稻根际土壤微生物对碳源的利用程度和碳源代谢的多样性差异显著, 处理后10 d, GN处理和G处理根际微生物对6类碳源的利用程度显著高于S处理, GN处理根际土壤微生物对氨基酸、酚酸、羧酸类碳源的利用程度显著高于G处理。GN处理和G处理的根系伤流量分别比S处理的提高27.27%和14.84%, 再生季分蘖数和产量分别比S处理的提高102.50%~111.11%、42.50%~44.44%和91.41%~108.72%、37.93%~40.94%。由此可见, 再生季采用干湿交替灌溉或施用促苗肥均可提高根际土壤酶活性, 促进再生季稻根际土壤微生物对碳源的利用程度和代谢多样性, 从而有利于根际土壤有机质的氧化和腐殖质的形成及增加根际土壤中养分的有效性, 促进再生季新根的形成和腋芽的萌发。采用干湿交替灌溉耦合促苗肥对再生季稻的促控效果是最佳的。

关键词: 再生稻; 水肥处理; 根际土壤酶活性; 根际微生物

Effects of Different Irrigation and Fertilizer Application Regimes on Soil Enzyme Activities and Microbial Functional Diversity in Rhizosphere of Ratooning Rice

CHEN Hong-Fei^1,^*, PANG Xiao-Min^1,^*, ZHANG Ren², ZHANG Zhi-Xing¹, XU Qian-Hua³, FANG Chang-Xun¹, LI Jing-Yong⁴, LIN Wen-Xiong^1,^*

¹ College of Life Sciences, Fujian Agriculture and Forestry University / Fujian Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou 350002, China

²Agriculture Bureau of Jianyang District, Nanping City, Fujian Province, Jianyang 353000, China

³Planting Technique Extension Station of Fujian Province Agriculture Department, Fuzhou 350001, China

⁴Chongqing Academy of Agricultural Sciences, Chongqing 402160, China

Fund:This study was supported by the National Key Research and Development Program of China (2016YFD0300508), the Natural Foundation of Fujian Higher Education Institutions for Young Scientists (Key Project) (JZ160435), and the Sci-tech Innovation Fund Project of Fujian Agriculture and Forestry University (CXZX2016076, CXZX2016077).

Abstract

Irrigation and fertilizer application regime is one of key measures for high and stable yield of ratooning rice, especially for middle or low stubble ratooning rice by machine-harvest. Ascertaining rational management of water and fertilization is important for improving the axillary bud germination rate of lower nodes in middle or low stubble ratooning rice by machine-harvest. With Luyoumingzhan as a test material, the effects of different irrigation and fertilizer application regimes on ratooning tillers, soil enzyme activity and rhizosphere microbial functional diversity of ratooning rice were studied by setting up three treatments: (1) dry-wet alternate irrigation and nitrogen application for ratooning bud development (GN); (2) dry-wet alternate irrigation and no nitrogen application for ratooning bud development (G); (3) flooding irrigation and no nitrogen application for ratooning bud development (S). The different irrigation and fertilization treatments had a significant impact on the rhizosphere Eh potential and rhizosphere soil enzymes. On the 10th day after treatment, the rhizosphere soil Eh levels and the activities of rhizosphere soil polyphenol oxidase, peroxidase, phosphate monoester enzyme and hydrogen peroxide enzyme were significantly improved in G than in S. The activities of rhizosphere soil polyphenol oxidase, peroxidase, urease, invertase, phosphate monoester enzyme and hydrogen peroxide enzyme were significantly improved in GN than in G. The rhizosphere soil microbe differed significantly in use of single carbon sources and diversity of carbon metabolism under different treatments. On the 10th day after treatment, the uses of single carbon sources and diversity of carbon metabolism in GN and G were significantly greater than those in S. The abilities to utilize amino acids, phenolic acids, carboxylic acid in GN were higher than those in G. Compared with S, bleeding rate of GN and G was increased by 27.27% and 14.84%, ratooning rice tiller and yield were increased by 102.50%-111.11%, 42.50%-44.44% and 91.41%-108.72%, 37.93%-40.94%, respectively. All these results indicated that dry-wet alternate irrigation or nitrogen application for ratooning bud development could improve rhizosphere soil enzyme activities and promote the uses of single carbon sources and diversity of carbon metabolism, which is conducive to increasing the effectiveness of nutrient, the formation of new roots and the axillary buds germination. The effects of dry-wet alternate irrigation coupling with fertilizer are the best for ratooning rice growth.

Keyword: Ratoon rice; Irrigation and fertilization treatments; Rhizosphere soil enzymes; Rhizosphere microorganisms

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再生稻是一种两收的资源节约型稻作方式, 已成为南方稻区稳粮增收、优化种植结构和提高种粮效益的重要稻作制度。近年来, 随着劳动力成本的不断上升, 机收再生稻逐步成为再生稻的主要种植模式, 采用中低留桩高度成为生产上解决机收再生稻碾压区与非碾压区再生季成熟期不一致和降低机收碾压导致再生季减产的主要措施^{[1, 2, 3]}, 随着留桩高度的降低, 再生季主体分蘖也由以高节位再生腋芽为主转变为以低节位再生腋芽为主^[4], 因此, 促进低节位再生腋芽萌发成穗是提高机收再生稻再生季产量、发挥再生稻稻作制度作用的关键。众多研究结果表明再生腋芽的萌发成穗与再生季稻根系活性有着显著或极显著的相关性^{[5, 6]}, 再生季稻根系机能对再生腋芽萌发的影响在中低留桩蓄留再生稻时尤为明显; 作物的根系形态和机能除了与品种特性有关外, 还受到环境(水分、营养条件等)的影响^[7]。近年来, 许多学者开展了再生季氮肥不同施用量、不同施用比例对机收中低留桩再生稻根系机能和腋芽萌发影响的研究^{[1, 8, 9, 10]}, 但关于再生季水分和氮肥耦合对机收中低留桩再生稻根系机能和腋芽萌发影响的研究却鲜有报道。已有研究表明, 根际是作物-土壤-微生物及其环境条件相互作用的界面, 也是各种养分、水分、有益和有害物质及生物作用于根系或进入根系参与循环的门户^[11], 因此, 要阐明再生季水分和氮肥耦合对再生稻根系机能和腋芽萌发的影响, 还必须从根际生态学的角度深入分析。本试验设置再生季不同水肥处理, 探讨对再生稻根系机能及根际土壤酶活性和微生物功能多样性的影响, 以期揭示水肥调控再生稻根系机能及腋芽萌发的根际生物学过程, 为机收中低留桩再生稻高产栽培和定向调控提供理论依据。

1 材料与方法

1.1 试验材料与设计

试验于2014— 2015年在福建农林大学教学基地(26° 10′ N, 119° 23′ E)进行, 供试土壤为沙壤土, 土壤pH 5.65, 含全氮1.98 g kg^-1、碱解氮127.90 mg kg^-1、有效磷43.10 mg kg^-1、速效钾137.51 mg kg^-1、有机质23.83 g kg^-1。

试验材料为福建省三明市农业科学院选育的杂交稻品种泸优明占, 采用盆栽种植的方式培养, 塑料桶高0.30 m, 顶部直径0.30 m, 底部直径0.23 m, 每桶移栽3株水稻, 头季于3月24日播种, 5月4日移栽, 7月18日齐穗, 8月21日收割, 留桩25 cm。头季收割后进行3种水肥处理(S, G, GN), S处理为再生季分蘖期实行淹水灌溉(始终保持水层2~4 cm), 不施用促苗肥; G处理为再生季分蘖期干湿交替灌溉(灌1~2 cm薄水层后自然落干, 直至土壤水势达到-15 kPa时复水至水层1~2 cm), 不施用促苗肥; GN处理为再生季分蘖期干湿交替灌溉(灌1~2 cm薄水层后自然落干, 直至土壤水势达到-15 kPa时复水至水层1~2 cm), 施用促苗肥, 在土壤中预埋Watermark Model 200型土壤水分传感器(Irrometer Corporation, 美国)监测水势, 每天下午测水势, 当数值达到限定值时复水, 3个处理再生季氮肥施用见表1, 每个处理40桶, 10月3日再生季齐穗, 11月5日再生季成熟, 3个处理除上述时期水肥管理措施不同外, 头季和再生季其他田间管理措施相同。

表1 再生季稻不同生育时期氮肥施用 Table 1 Nitrogen application at different growth stages in ratooning rice

1.2 土壤取样

于头季稻收割后水肥处理前(before irrigation and fertilizer treatments, BIFT)和水肥处理后10 d (10 d after irrigation and fertilizer treatments, AIFT)取土壤样品, 每次每个处理随机选取6株盆栽稻株, 用灭菌过的剪刀剥除离根系较远的土壤, 露出根际土, 再尽量取紧靠根面的土壤, 用剪刀轻轻剥掉根际土, 混匀后采用五点取样法取土样, 分为2份分别保存于4℃冰箱和-80℃冰箱备用。

1.3 根系伤流量的测定

参照李合生贴棉吸附法测定伤流量^[12], 分别在水肥处理前和处理后10 d测定每个处理6株伤流量, 于傍晚7点到次日早上7点收集伤流, 称重, 计算伤流强度。

1.4 再生稻分蘖数观察

头季稻收割后, 选取和稻桩平均数一样或相近的水稻稻桩挂牌, 选取每个处理9株稻桩挂牌标记, 每隔5 d观察再生分蘖数。

1.5 产量和产量构成因子测定

再生季收割时, 选取每处理6株考察有效穗、穗粒数、结实率和千粒重。

1.6 土壤酶活性测定

参照关松荫等^[13]、鲁如坤^[14]方法测定土壤酶活性, 采用苯酚钠比色法测定土壤脲酶活性; 采用3, 5-二硝基水杨酸比色法测定蔗糖酶活性; 采用高锰酸钾滴定法测定过氧化氢酶活性; 采用对硝基苯磷酸盐法测定磷酸单酯酶活性; 采用邻苯三酚比色法测定多酚氧化酶、过氧化物酶活性。

1.7 土壤微生物群落功能多样性测试

采用BIOLOG ECO微平板法, 称取5 g鲜土于灭菌的三角瓶中, 加入100 mL灭菌生理盐水, 封口, 120 r min^-1振荡30 min, 冰上静置2~5 min, 于超净台中轻轻取上清液5 mL移于灭菌的100 mL三角瓶中, 并且加入45 mL无菌水, 摇匀得到1∶ 100提取液。重复稀释3次, 制得1∶ 1000的提取液, 立即用于ELISA反应。将BIOLOG ECO平板预热到25℃, 用移液枪吸取1∶ 1000的提取液150 μ L于各个孔中, 其中A₁、A₅、A₉为150 μ L无菌水做空白对照。28℃恒温培养1周, 在0、1、2、3、4、5、6、7 d用ELISA反应平板读数器读取OD_590nm的吸光值。

1.8 数据分析处理

因2年间试验处理的趋势基本一致, 本文仅采用2015年数据分析讨论; 微生物群落结构用AWCD作为整体活性的有效指数, AWCD = Σ (C-R)/31, 式中C为每孔的吸光度, R为对照孔的吸光值; 选择BIOLOG ECO微平板法培养96 h的数据计算Simpson优势度指数(J)、Shannon-Wienner多样性指数(H)和Shannon均匀度指数(E), J= 1-Σ P_i², H= -Σ (P_i· ln P_i), E=H/ln S, 式中P_i为第i孔相对吸光度值与整个平板相对吸光度值总和的比值, S为发生颜色变化的孔数; 采用Windows 4.5软件作主成分分析(Principal Component Analysis, PCA), R2.14.0软件作聚类分析; 用Microsoft Excel 2013软件进行数据计算和作图, 用DPS 7.05软件进行差异性分析。

2 结果与分析

2.1 不同水肥处理对再生季稻分蘖动态及产量构成的影响

从图1可知, 再生分蘖增长呈先快后慢趋势, 水肥处理后5 d是再生分蘖的快速增长期, 5~15 d再生分蘖进入缓慢增长期, 15 d后基本达到稳定, 但是不同水肥处理再生分蘖的增长速度不同, 水肥处理后5 d, G处理的再生分蘖增长略高于S处理, GN处理的再生分蘖增长明显高于G处理和S处理, 水肥处理后5~15 d, G处理增长速度加快, GN处理和G处理均高于S处理, 至水肥处理后25 d, GN处理的再生分蘖分别比G处理、S处理多46.15%和72.73%。可见, 不同水肥运筹与再生分蘖有着密切联系, 水肥耦合可明显促进休眠芽萌发成再生分蘖。

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图1 再生稻不同水肥调控下再生分蘖情况 GN: 干湿交替灌溉施用促苗肥处理; G: 干湿交替灌溉不施用促苗肥处理; S: 淹水灌溉不施用促苗肥处理。Fig. 1 Tiller amounts of ratooning rice under different irrigation and fertilizer treatments GN: dry-wet alternate irrigation with nitrogen application for bud development; G: dry-wet alternate irrigation without nitrogen application for bud development; S: flood irrigation without nitrogen application for bud development.

进一步从表2可知, 与S处理的再生季产量相比, GN处理和G处理有显著的增产作用, GN处理和G处理的再生季产量分别比S处理的提高91.41%~ 108.72%和37.93%~40.94%。在再生季稻产量构成因子中, 3个处理间千粒重和每穗粒数基本相近, 差异未达显著水平, 结实率以S处理最高, 显著高于GN处理和G处理, GN处理和G处理间的结实率差异不显著; 单位面积有效穗数以GN处理最高, G处理次之, GN处理、G处理分别比S处理高102.50%~ 111.11%和42.50%~44.44%, 3个处理间的差异达显著水平, 表明再生季合理的水分管理和肥料措施确能促进再生腋芽的萌发, 从而通过提高再生季单位面积有效穗实现增产, 但是水肥耦合在促进休眠腋芽萌发的同时, 结实率却有所下降, 这主要是低节位再生分蘖结实率相对较低导致的。

表2 不同水肥处理下再生季水稻盆栽产量及其构成因素 Table 2 Yield and its components of ratooning rice in pot experiment under different irrigation and fertilizer treatments

2.2 不同水肥处理对再生季稻根系伤流强度的影响

根系伤流量的变化可以间接地反映水稻的根系活力情况。再生稻不同水肥处理之间的根系伤流量存在显著差异(表3), 与处理前相比, 3个处理的再生季根系伤流量均有所提高, 但提高幅度不同, 再生季水肥处理后10 d, GN处理的根系伤流量为116.72 g m^-2 h^-1, G处理的根系伤流量为105.34 g m^-2 h^-1, GN处理比G处理提高10.82%, GN处理和G处理分别比S处理(91.71 g m^-2 h^-1)提高27.27%和14.84%, 差异达显著水平, 表明再生季合理的水分管理或肥料措施均对根系特性有显著影响, 且水肥耦合促进再生季稻根系特性的效应较单一措施更为显著。

表3 不同水肥处理下再生季稻根系伤流强度的变化 Table 3 Bleeding rate of ratooning crop under different irrigation and fertilizer treatments (g m^-2 h^-1)

2.3 不同水肥处理对再生季水稻根际土壤酶活性的影响

土壤酶参与土壤中各种生物化学过程, 其活性的高低直接反映土壤生物化学过程的强弱^[15], 再生季稻不同水肥处理的土壤酶活性差异显著(图2), 与水肥处理前相比, GN处理可以显著提高再生季稻根际土壤的多酚氧化酶、过氧化物酶、脲酶、蔗糖酶、磷酸单酯酶、过氧化氢酶活性, G处理可以显著提高多酚氧化酶、磷酸单酯酶、过氧化氢酶的活性, S处理可以显著提高过氧化氢酶的活性, 而多酚氧化酶、过氧化物酶、磷酸单酯酶活性显著降低; 处理后10 d, 与S处理相比, G处理的土壤多酚氧化酶、过氧化物酶、磷酸单酯酶、过氧化氢酶活性显著提高, 与G处理相比, GN处理的根际土壤多酚氧化酶、过氧化物酶、脲酶、蔗糖酶、磷酸单酯酶、过氧化氢酶活性显著提高。

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图2 再生季水肥调控对根际土壤酶活性的影响图柱上不同小写字母表示处理间在0.05水平上差异显著。缩写同图1。Fig. 2 Effect of different irrigation and fertilizer treatments on rhizospheric soil enzyme activities in ratooning rice
Bars indexed with different small letters are significantly different among treatments at the 0.05 probability level. Abbreviations are the same as those given in Figure 1.

2.4 不同水肥处理对土壤Eh和pH值的影响

Eh是反映土壤氧化还原的重要指标, 土壤的Eh适宜范围是200~400 mV。从图3可知, 水肥处理前后, 3个处理的Eh均有所提高, 但提高幅度不同, 处理后10 d, S处理的土壤Eh虽有所提高但未达显著水平, GN处理和G处理的土壤Eh显著高于处理前, 与S处理的差异也达显著水平。在头季稻收割后土壤的pH值显著提高, 但各个处理之间差异不显著。可见, GN处理和G处理可以提高土壤的通透性和氧含量, 从而显著提高土壤Eh, 但对土壤pH值并没有显著影响。

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图3 不同水肥调控对再生季水稻土壤Eh、pH值的影响图柱上不同小写字母表示处理间在0.05水平上差异显著。缩写同图1。Fig. 3 Effect of different irrigation and fertilizer treatments on rhizospheric soil Eh and pH values in ratooning rice Bars indexed with different small letters are significantly different among treatments at the 0.05 probability level. Abbreviations are the same as those given in Figure 1.

2.5 不同水肥处理再生季水稻根际土壤微生物的Biolog分析

2.5.1 再生季水稻根际土壤微生物对碳源利用程度的动力学特征从图4可以发现, 各处理根际微生物对碳源的利用程度随着培养时间的延长而增加, 但存在较大差异, 处理后10 d的土壤AWCD值为GN处理> G处理> S处理> 处理前, 表明再生季干湿交替灌溉耦合促苗肥处理明显促进了根际土壤微生物群落对碳源的利用程度。

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图4 再生季水肥处理前后根际土壤所有碳源平均颜色(AWCD值)随时间的变化情况 GN10: 干湿交替灌溉施用促苗肥处理后10 d; G10: 干湿交替灌溉不施用促苗肥处理后10 d; S10: 淹水灌溉不施用促苗肥处理后10 d; BIFT: 水肥处理前。Fig. 4 Average well colour development (AWCD) of rhizospheric soil before and after treating GN10: 10 d after dry-wet alternate irrigation with nitrogen application for bud development; G10: 10 d after dry-wet alternate irrigation without nitrogen application for bud development; S10: 10 d after flood irrigation without nitrogen application for bud development. BIFT: before irrigation and fertilizer treatments.

2.5.2 再生季水稻根际土壤微生物群落多样性和均匀度指数分析土壤微生物群落多样性是群落总体动态变化的反映, 从表4可知, 处理后10 d, GN处理和G处理根际微生物群落多样性和均匀度指数均显著高于S处理和处理前, GN处理和G处理之间的差异也达显著水平, S处理和处理前的差异不显著, 可见, 再生季合理的水分管理有助于再生稻根际土壤细菌群落多样性和均匀度的提高, 合理的水分管理耦合施用促苗氮肥则效果更显著。

表4 再生季不同水肥运筹方式下根际土壤细菌群落多样性和均匀度指数分析 Table 4 Diversity and evenness analysis of rhizospheric soil bacterial communities under different irrigation and fertilizer treatments in ratooning rice

2.5.3 再生季水稻根际土壤微生物对6类碳源的利用程度对土壤微生物结构的研究, 可以从土壤微生物对碳源利用程度的差异着手。图5表明再生季不同水肥调控下根际土壤微生物对不同类型碳源的利用程度差异显著, 处理后10 d, GN处理和G处理根际微生物对6类碳源的利用程度显著高于S处

理, GN处理根际土壤微生物对氨基酸、酚酸、羧酸类碳源的利用程度显著高于G处理, 而对胺类、聚合物类和碳水化合物类3类碳源的利用程度GN处理和G处理相近, 差异不显著。由此可见, 再生季合理的水分管理可以显著提高根际微生物对6类碳源的利用程度, 在此基础上, 适当施用促苗氮肥可以显著提高根际微生物对部分类型碳源的利用程度。

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图5 不同水肥处理下根际土壤微生物对不同类型碳源的代谢利用 GN10: 干湿交替灌溉施用促苗肥处理后10 d; G10: 干湿交替灌溉不施用促苗肥处理后10 d; S10: 淹水灌溉不施用促苗肥处理后10 d; BIFT: 水肥处理前。Fig. 5 Effect of different irrigation and fertilizer treatments on utilization of different sources of carbon in rhizospheric soil microbes GN10: 10 d after dry-wet alternate irrigation with nitrogen application for bud development; G10: 10 d after dry-wet alternate irrigation without nitrogen application for bud development; S10: 10 d after flood irrigation without nitrogen application for bud development. BIFT: before irrigation and fertilizer treatments.

2.5.4 再生季水稻根际土壤微生物对碳源底物利用差异的主成分及聚类分析从图6-A可知, 培养96 h的AWCD值能够较好地区分不同水肥处理再生季水稻根际土壤微生物群落对于碳源底物的代谢利用差异。在不同水肥处理后, 主成分1 (PC1)和主成分2 (PC2)分别解释变量方差的49.7%和38.4%, 与主成分1 (PC1)正相关系数最高的单一碳源是Tween 40 (polymers)、D-甘露醇(carbohydrates); 与主成分2 (PC2)负相关系数绝对值最高的单一碳源为α -环式糊精(polymers)和苯乙胺(amines amides)。

图6-B表明, 不同水肥处理下再生季水稻根际微生物碳源代谢之间存在明显差异。GN和G处理比处理前和S处理大, 施肥处理比未施肥处理大; G处理和GN处理聚成一类, S处理和处理前聚为一类, 并且各重复样品之间的相关性较好。不同水肥处理再生季水稻的BIOLOG数据分析表明, G处理根际土壤微生物群落的碳源代谢多样性大于S处理和处理前, 在此基础上, 施用促苗肥会进一步提高再生季水稻根际土壤微生物群落碳源代谢的多样性。

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图6 BIOLOG主成分分析图(A)和聚类分析图(B) (A) 1~3: GN处理; 4~6: G处理; 7~9: S处理; 10~12: 水肥处理前。(B) GN10.1, GN10.2, GN10.3: 干湿交替施用促苗肥处理后10 d; G10.1, G10.2, G10.3: 干湿交替不施用促苗肥处理后10 d; S10.1, S10.2, S10.3: 淹水不施用促苗肥处理后10 d; X1, X2, X3: 水肥处理前; A1~A31: 31种碳源。Fig. 6 Classification of treatments by principal component (A) and cluster (B) analysis with BIOLOG data (A) 1-3: GN treatment; 4-6: G treatment; 7-9: S treatment; 10-12: before irrigation and fertilizer treatments. (B) GN10.1, GN10.2, GN10.3: 10 d after dry-wet alternate irrigation with nitrogen application for bud development; G10.1, G10.2, G10.3: 10 d after dry-wet alternate irrigation without nitrogen application for bud development; S10.1, S10.2, S10.3: 10 d after flood irrigation without nitrogen application for bud development; X1, X2, X3: before irrigation and fertilizer treatments. A1-A31: thirty-one kinds of carbon sources.

3 讨论

再生季稻产量与头季收割后稻桩上休眠腋芽萌发成穗数密切相关^[16]。目前, 随着我国经济的发展和农村劳动力的大量转移, 导致季节性缺劳动力, 劳动力价格急剧攀升, 机械化轻简再生稻栽培成为再生稻规模化种植发展的必然选择; 机械化轻简再生稻栽培一般采用中低留桩高度来解决机收再生稻碾压区与非碾压区再生季成熟期的不一致和降低机收碾压导致的再生季减产, 因此, 机收中低留桩再生稻低节位腋芽的促发栽培成为机械化轻简再生稻高产稳产栽培的关键; 大量的研究结果表明再生稻休眠芽尤其是低节位腋芽的萌发生长成穗与再生季水稻根系活性有着显著或极显著的相关性^{[6, 17]}。再生

稻根系由两部分组成, 一是母茎上存活的老根与其在再生季产生的新根, 二是稻桩母茎茎节休眠芽上根原基萌发而成的新根^[16], 其中再生新根对机械化收割中低留桩蓄留再生稻再生分蘖的萌发尤为重要, 两者间存在着“ 根芽同伸” 关系。作物的根系形态和机能除了与品种特性有关外, 还受到环境(水分、营养条件)的影响^[7]。本研究发现再生季单一的干湿交替灌溉或施用促苗肥均能显著提高再生季根系活性, 而水肥耦合促进再生季稻根系活性的效应较单一措施则更为显著, 因而, GN处理的再生分蘖分别比G处理、S处理多46.15%和72.73%, 表明再生季合理的水分管理和肥料措施耦合确能更为显著地提高再生季根系活性进而促进再生腋芽的萌发, 提高再生季单位面积有效穗实现增产, 而且增产主要是通过促进低节位腋芽的萌发成穗来实现的。这与俞道标等的研究结果一致^[18], 也表明了再生季水、肥运筹通过调控再生季水稻的根系活性进而影响再生腋芽的萌发和再生季稻产量。

环境条件是通过何种途径调控作物的根系构型呢?张福锁等^[19]认为栽培调控的实质是调控根际环境, 为作物创造一个良好的根际生态环境从而调控作物-土壤-微生物间的相互作用达到促进作物生长和发育的目的, 根际是作物-土壤-微生物及环境条件相互作用的区域, 也是各种营养物质、水分、有益和有害物质及生物作用于根系或进入根系参与循环的界面。作物-土壤-微生物及其环境之间的相互作用构成了作物的根际生态系统, 它对于土壤的理化性质、根系的形态、生理有着重要的影响^{[20, 21]}, 根际化学过程包括根际土壤的酸碱反应、氧化还原反应等, 根际土壤酶和根际微生物是根际生态系统中重要组成部分, 在根际生态中起着重要的作用, 它们对于植物根际生态系统的物质循环和能量流动具有重要的影响, 由于根际土壤酶和根际微生物对外界条件尤其是土地管理、耕作制度的影响异常敏感, 其变化和群落组成、多样性常常被用作表征土壤质量变化的指标^[22]。土壤酶中, 脲酶能分解有机物, 促其水解成氨和二氧化碳; 多酚氧化酶与土壤有机质的形成有关, 是腐殖化的一种媒介; 过氧化物酶是土壤中一种重要的氧化还原酶类, 在有机质氧化和腐殖质形成过程中缺少不了过氧化物酶的作用; 蔗糖酶与土壤肥力的改善和提高关系密切, 它的主要作用是利于土壤中有机质的转化; 磷酸酶对于土壤有机磷化合物的转化有重要的作用, 土壤中有机磷

的分解强度取决于其活性高低的变化; 过氧化氢酶活性对于土壤腐殖质化强度大小和有机质积累程度有表征作用。本研究发现再生季稻不同水肥处理的根际氧化还原电位和根际土壤酶活性具有显著差异, 处理后10 d, GN处理和G处理可以提高根际土壤的氧化还原电位, 但对根际土壤pH值的变化没有显著影响, 与S处理相比, G处理对于根际土壤中的多酚氧化酶、过氧化物酶、磷酸单酯酶、过氧化氢酶活性有显著提高作用。Waldrop等^[23]的研究结果也指出土壤多酚氧化酶和过氧化物酶的活性对于土壤水分的变化比较敏感, 水分的增加会降低这两种酶的活性, 与G处理相比, GN处理对于六大土壤酶活性都有显著提高作用, 这与周卫等^[24]和赵之重^[25]的报道相似, 表明再生季干湿交替灌溉有利于提高土壤的通透性和氧含量, 对于根际土壤中有机质的氧化和腐殖质的形成有促进作用, 而水肥耦合措施对于根际土壤有机质的氧化和腐殖质的形成及增加根际土壤中养分有效性的促进作用要显著高于单一措施, 从而为再生季新根的形成和腋芽的萌发提供相应的物质需求保障。

根际土壤微生物是根际土壤酶的重要来源, 同时也直接参与根际的生物过程, 影响根系构型的形成和特性。基于BIOLOG平板法的根际微生物功能多样性研究表明不同水肥调控下再生季稻根际土壤微生物对碳源的利用程度和碳源代谢的多样性差异显著, 再生季干湿交替水分管理下根际土壤微生物群落对碳源的利用程度和碳源代谢的多样性显著大于淹水处理和处理前。在合理水分管理的基础上, 适当施用促苗肥有利于进一步提高根际微生物对氨基酸、酚酸、羧酸类碳源的利用程度和碳源代谢的多样性。从31个碳源因子中提取与碳源利用相关的主成分1和主成分2能够解释变量方差总和的88.1%, 在两大成分分离中起主要贡献作用的是聚合物类、碳水化合物类和氨基酸类, 土壤碳水化合物类和氨基酸类是土壤中微生物细胞必需的组成物质和活动的主要能源, 不同水肥调控下根际土壤微生物对根系环境中碳水化合物和氨基酸的利用程度不同, 直接影响根际土壤微生物的数量和种类, 根际土壤微生物数量和种类的不同变化又反过来影响着作物根系的生长发育^{[26, 27]}。可见, 再生季合理水分管理和适当促苗肥耦合引起的再生季稻根际土壤微生物对碳源利用程度和碳源代谢多样性的提高, 是导致GN处理下再生季根际土壤养分有效性和根系活性增强的重要原因。

4 结论

不同水肥调控措施对机收再生稻再生季根际土壤酶活性及微生物功能多样性有显著影响, 与淹水灌溉不施用促苗肥和干湿交替灌溉不施用促苗肥相比, 再生季干湿交替灌溉且施用促苗肥能显著提高根际土壤的氧化还原电位和根际土壤酶活性, 提高根际土壤微生物群落对碳源的利用程度和碳源代谢的多样性, 从而提高机收再生稻再生季的根系机能和产量。因而, 再生季分蘖期采用干湿交替灌溉耦合施用促苗肥是机收中低留桩再生稻高产高效栽培的关键技术之一。

致谢: 本研究得到闽台作物特色种质创制与绿色栽培协同创新中心[闽教科(2015)75号]提供的技术帮助, 特此致谢。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

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[11]	李春俭, 马玮, 张福锁. 根际对话及其对植物生长的影响. 植物营养与肥料学报, 2008, 14: 178-183 Li C J, Ma W, Zhang F S. Rhizosphere talk and its impacts on plant growth. Plant Nutr Fert Sci, 2008, 14: 178-183 (in Chinese with English abstract) [本文引用:1]
[12]	李合生. 植物生理生化实验原理和技术. 北京: 高等教育出版社, 2000. pp 164-261 Li H S. Experiment Principle and Technique on Plant Physiology and Biochemistry. Beijing: Higher Education Press, 2000. pp 164-261(in Chinese) [本文引用:1]
[13]	关松荫, 张德生, 张志明. 土壤酶及其研究法. 北京: 农业出版社, 1986. pp 274-325 Guan S Y, Zhang D S, Zhang Z M. Soil Enzymes and Its Research Methods. Beijing: Agriculture Press, 1986. pp 274-325(in Chinese) [本文引用:1]
[14]	鲁如坤. 土壤农业化学分析方法. 北京: 中国农业科技出版社. 2000. pp 252-254 Lu R K. Analysis Method of Soil Agricultural Chemistry. Beijing: China Agricultural Science and Technology Press, 2000. pp 252-254(in Chinese) [本文引用:1]
[15]	万忠梅, 吴景贵. 土壤酶活性影响因子研究进展. 西北农林科技大学学报(自然科学版), 2005, 33(6): 87-92 Wan Z M, Wu J G. Study progress on factors affecting soil enzyme activity. J Northwest Sci-Tech Univ Agric & For(Nat Sci Edn) , 2005, 33(6): 87-92 (in Chinese with English abstract) [本文引用:1]
[16]	林文雄, 陈鸿飞, 张志兴, 徐倩华, 屠乃美, 方长旬, 任万军. 再生稻产量形成的生理生态特性与关键栽培技术的研究与展望. 中国生态农业学报, 2015, 23(4): 392-401 Lin W X, Chen H F, Zhang Z X, Xu Q H, Tu N M, Fang C X, Ren W J. Research and prospect on physio-ecological properties of ratoon rice yield formation and its key cultivation technology. Chin J Eco-Agric, 2015, 23(4): 392-401 (in Chinese with English abstract) [本文引用:2]
[17]	林文, 李义珍, 姜照伟, 郑景生. 再生稻根系形态和机能的品种间差异及与产量的关联性. 福建农业学报, 2001, (1): 1-4 Lin W, Li Y Z, Jiang Z W, Zheng J S. Morphologic and functional difference of root systems among ratooning rice varieties and its correlation with yield. Fujian J Agric Sci,* 2001*, (1): 1-4 (in Chinese with English abstract) [本文引用:1]
[18]	俞道标, 赵雅静, 黄顽春, 李小萍, 姜照伟, 陈双龙. 低桩机割再生稻生育特性和氮肥施用技术研究. 福建农业学报, 2012, 27: 485-490 Yu D B, Zhao Y J, Huang W C, Li X P, Jiang Z W, Chen S L. Growth characteristics and nitrogen application technique of ratoon rice with machine harvest in low cutting. Fujian J Agric Sci, 2012, 27: 485-490 (in Chinese with English abstract) [本文引用:1]
[19]	张福锁, 王激清, 张卫峰, 崔振岭, 马文奇, 陈新平, 江荣风. 中国主要粮食作物肥料利用率现状与提高途径. 土壤学报, 2008, 45: 915-924 Zhang F S, Wang J Q, Zhang W F, Cui Z L, Ma W Q, Chen X P, Jiang R F. Nutrient use efficiencies of major cereal crops in China and measures for improvement. Acta Pedol Sin, 2008, 45: 915-924 (in Chinese with English abstract) [本文引用:1]
[20]	沈建凯, 黄璜, 傅志强, 张亚, 陈灿, 金小马, 廖晓兰, 龙攀, 高文娟, 谢小婷. 规模化稻鸭生态种养对稻田杂草群落组成及物种多样性的影响. 中国生态农业学报, 2010, 18(1): 123-128 Shen J K, Huang H, Fu Z Q, Zhang Y, Chen C, Jin X M, Liao X L, Long P, Gao W J, Xie X T. Effect of large-scale rice-duck eco-farming on the composition and diversity of weed community in paddy fields. Chin J Eco-Agric, 2010, 18(1): 123-128 (in Chinese with English abstract) [本文引用:1]
[21]	裴鹏刚, 张均华, 朱练峰, 禹盛苗, 金千瑜. 根际氧浓度调控水稻根系形态和生理特性研究进展. 中国稻米, 2013, 19(2): 6-8 Pei P G, Zhang J H, Zhu L F, Yu S M, Jin Q Y. Research progress on effects of rhizosphere dissolved oxygen on root morphology and physiological characteristics in rice. China Rice, 2013, 19(2): 6-8 (in Chinese) [本文引用:1]
[22]	胡婵娟, 郭雷, 刘国华. 黄土丘陵沟壑区不同植被恢复格局下土壤微生物群落结构. 生态学报, 2014, 34: 2986-2995 Hu C J, Guo L, Liu G H. Soil microbial community structure under different vegetation restoration patterns in the loess hilly area. Acta Ecol Sin, 2014, 34: 2986-2995 (in Chinese with English abstract) [本文引用:1]
[23]	Waldrop M P, Zak D R, Sinsabaugh R L, Gallo M, Lauber C. Nitrogen deposition modifies soil carbon storage through changes in microbial enzymatic activity. Ecol Appl, 2004, 14: 1172-1177 [本文引用:1]
[24]	周卫, 刘经荣, 张德远. 水稻土有机-无机肥料配合施用的效应(之四). 江西农业大学学报, 1990, (2): 24-31 Zhou W, Liu J R, Zhang D Y. The effect of mixed application of organic and inorganic fertilizers in paddy soil (the fourth report). Acta Agric Univ Jiangxiensis, 1990, (2): 24-31 (in Chinese with English abstract) [本文引用:1]
[25]	赵之重. 土壤酶与土壤肥力关系的研究. 青海大学学报(自然科学版), 1998, (3): 24-29 Zhao Z Z. The Relationships between soil enzyme and soil fertility. J Qinghai Univ, 1998, (3): 24-29 (in Chinese with English abstract) [本文引用:1]
[26]	高子勤, 张淑香. 连作障碍与根际微生态研究: I. 根系分泌物及其生态效应. 应用生态学报, 1998, 9: 549-554 Gao Z Q, Zhang S X. Continuous cropping obstacle and rhizospheric microecology: I. Root exudates and their ecological effects. Chin J Appl Ecol, 1998, 9: 549-554 (in Chinese with English abstract) [本文引用:1]
[27]	张淑香, 高子勤. 连作障碍与根际微生态研究: II. 根系分泌物与酚酸物质. 应用生态学报, 2000, 11: 153-157 Zhang S X, Gao Z Q. Continuous cropping obstacle and rhizospheric microecology: II. Root exudates and phenolic acids. Chin J Appl Ecol, 2000, 11: 153-157 (in Chinese with English abstract) [本文引用:1]

2013

0.0

... 近年来, 随着劳动力成本的不断上升, 机收再生稻逐步成为再生稻的主要种植模式, 采用中低留桩高度成为生产上解决机收再生稻碾压区与非碾压区再生季成熟期不一致和降低机收碾压导致再生季减产的主要措施^[1,2,3], 随着留桩高度的降低, 再生季主体分蘖也由以高节位再生腋芽为主转变为以低节位再生腋芽为主^[4], 因此, 促进低节位再生腋芽萌发成穗是提高机收再生稻再生季产量、发挥再生稻稻作制度作用的关键 ...

... 近年来, 许多学者开展了再生季氮肥不同施用量、不同施用比例对机收中低留桩再生稻根系机能和腋芽萌发影响的研究^[1,8,9,10], 但关于再生季水分和氮肥耦合对机收中低留桩再生稻根系机能和腋芽萌发影响的研究却鲜有报道 ...

2014

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2015

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2012

0.0

2010

0.0

... 众多研究结果表明再生腋芽的萌发成穗与再生季稻根系活性有着显著或极显著的相关性^[5,6], 再生季稻根系机能对再生腋芽萌发的影响在中低留桩蓄留再生稻时尤为明显 ...

2004

0.0

... 大量的研究结果表明再生稻休眠芽尤其是低节位腋芽的萌发生长成穗与再生季水稻根系活性有着显著或极显著的相关性^[6,17] ...

1996

0.0

... 作物的根系形态和机能除了与品种特性有关外, 还受到环境(水分、营养条件等)的影响^[7] ...

... 作物的根系形态和机能除了与品种特性有关外, 还受到环境(水分、营养条件)的影响^[7] ...

2013

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2010

0.0

2013

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2008

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李春俭, 马玮, 张福锁. 根际对话及其对植物生长的影响. 植物营养与肥料学报, 2008, 14: 178-183

Li C

, Ma

, Zhang F

. Rhizosphere talk and its impacts on plant growth. Plant Nutr Fert Sci, 2008, 14: 178-183 (in Chinese with English abstract)

The rhizosphere talk is the interactions between root and invading root systems of neighboring plants as well as root and soil-borne organisms including bacterium, fungi and animals in the rhizosphere. The exchanges of substances and energy and transmission of signals are involved in the rhizosphere talk. Increasing evidence suggests that root exudates might play an important role in the communication between root-root and root-soil organisms. These root-based interactions between plants and organisms in the rhizosphere are essentially the competition for limited space, water and mineral nutrients in the soil. In comparison with the study on the single process occurred in the rhizosphere, the conscious researches on rhizosphere talk and its impacts on plant growth are just started in the recent years. The progress of these studies is currently limited by the lack of the efficient and useful methods and techniques. In this paper, new results and trends of the researches on rhizosphere talk and its impacts on plant growth are introduced.

Department of Plant Nutrition, China Agricultural University，Key Laboratory of Plant Nutrient and Nutrient Cycle，Ministry of Agriculture, Key Laboratory of Plant-Soil Interactions, Ministry of Education, Beijing 100094, China

有了根际，也就有了根际对话。根际对话是发生在根际土壤中各种生物间，包括植物根系之间以及根系与土壤生物（细菌、真菌和土壤动物）之间的相互作用－即根际生物间的物质和能量交换以及信息传递。根系分泌物在根际对话中起着“语言”的作用。在根际对话中，植物起主导作用，对话的结果会对植物生长产生影响。根际生物之间相互作用的实质是对生长空间、水分和养分等资源的竞争。与根际研究相比，有意识地研究根际对话及其对植物生长影响才刚刚开始。研究方法和手段的局限性成为揭示根际对话机制的瓶颈。本文综述近年来国际上在根际对话及其对植物生长影响方面的研究进展和动态，期望我国在根际对话的基础和应用研究领域能够得到快速发展。

... 已有研究表明, 根际是作物-土壤-微生物及其环境条件相互作用的界面, 也是各种养分、水分、有益和有害物质及生物作用于根系或进入根系参与循环的门户^[11], 因此, 要阐明再生季水分和氮肥耦合对再生稻根系机能和腋芽萌发的影响, 还必须从根际生态学的角度深入分析 ...

2000

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... 3 根系伤流量的测定参照李合生贴棉吸附法测定伤流量^[12], 分别在水肥处理前和处理后10 d测定每个处理6株伤流量, 于傍晚7点到次日早上7点收集伤流, 称重, 计算伤流强度 ...

1986

0.0

... 6 土壤酶活性测定参照关松荫等^[13]、鲁如坤^[14]方法测定土壤酶活性, 采用苯酚钠比色法测定土壤脲酶活性 ...

2000

0.0

... 6 土壤酶活性测定参照关松荫等^[13]、鲁如坤^[14]方法测定土壤酶活性, 采用苯酚钠比色法测定土壤脲酶活性 ...

2005

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... 3 不同水肥处理对再生季水稻根际土壤酶活性的影响土壤酶参与土壤中各种生物化学过程, 其活性的高低直接反映土壤生物化学过程的强弱^[15], 再生季稻不同水肥处理的土壤酶活性差异显著(图2), 与水肥处理前相比, GN处理可以显著提高再生季稻根际土壤的多酚氧化酶、过氧化物酶、脲酶、蔗糖酶、磷酸单酯酶、过氧化氢酶活性, G处理可以显著提高多酚氧化酶、磷酸单酯酶、过氧化氢酶的活性, S处理可以显著提高过氧化氢酶的活性, 而多酚氧化酶、过氧化物酶、磷酸单酯酶活性显著降低 ...

2015

0.0

林文雄, 陈鸿飞, 张志兴, 徐倩华, 屠乃美, 方长旬, 任万军. 再生稻产量形成的生理生态特性与关键栽培技术的研究与展望. 中国生态农业学报, 2015, 23(4): 392-401

Lin W

, Chen H

, Zhang Z

, Xu Q

, Tu N

, Fang C

, Ren W

. Research and prospect on physio-ecological properties of ratoon rice yield formation and its key cultivation technology. Chin J Eco-Agric, 2015, 23(4): 392-401 (in Chinese with English abstract)

This article reviewed the current situations of ratoon rice in China and summarized the research progresses on the ratoon rice yield formation from the perspectives of the developmental genetics and physiological ecology. The key cultivation technology and eco-physiological mechanisms of the ratooning plants from low and high rice stubbles were comprehensively reviewed also. The ratoon rice technique gives two yields from one seeding, with significant yield advantages over single cropping rice and double cropping rice. Appropriate rice cultivars and cultivation methods are decisive factors of high yield of ratoon rice. For the ratoon rice from the high stubble of the hand-harvested first-season rice, to get high yield, the hybrid indica rice cultivars with heavy panicles were the best choices, which were relative weak in tillering ability for the first-season, and high regeneration capacity from lower nodes for the ratoon season. For the ratoon rice from the lower stubble of the mechanically harvested first-season rice, to get high yield, the hybrid indica rice with high regeneration capacity from lower nodes, or hybrid indica and japonica/indica rice crosses with the heavy panicle but weak photosensitivity were the best choices. Previous studies also suggested that roots adjustment was crucial to realize yield potential of ratoon rice. The ratoon rice root system was composed of the roots of the first-season rice and new roots of the ratoon season rice. The higher root activity from the mature stage of the first-season rice to the heading stage of the ratoon rice improved germination of axillary bud in rice stubbles and increased the panicles number of ratoon rice, which was favorable for high yield of ratoon rice. According to our study results and previous reports, we summarize the high-yielding cultural techniques of ratoon rice from hand-harvested first-season rice, such as early sowing in optimal time, ridge transplanting and furrow irrigation, two-time soil drying (heavy drought at tillering stage and moderate drought at 25 days after full heading stage in the first season), increased N application for bud development, and higher stubbles of 40 50 cm. In addition, the key cultivation technology and eco-physiological mechanisms of the ratoon rice from the low stubbles of machine-harvested first-season rice were comprehensively discussed from variety selection, fertilizer application, and the height of the remained stubbles. At last, the problems and future trends of ratoon rice were discussed. It was suggested that ratoon rice from low stubbles of mechanically harvested rice were the direction of ratoon rice development. The corresponding researches were therefore prospected.

本文介绍了目前我国再生稻的发展现状, 从发育遗传和生理生态角度综述了近年来再生稻产量形成的研究进展, 提出采用人工收割高留桩栽培再生稻时, 选择头季分蘖力相对较弱、再生季再生力强的重穗型杂交籼稻品种(组合)易获高产; 而采用机械化收割低留桩栽培再生稻时, 选择具强低位芽再生力的杂交籼稻品种或感光性弱的重穗型杂交粳稻品种(组合)、籼粳交水稻品种(组合)易获高产。头季成熟期至再生季齐穗期根系活力的强弱, 直接影响再生季产量的高低, 再生季稻高产的前提是健壮的头季根系和一定数量新生根系的有效结合, 这有利于促进腋芽的萌发成苗、增加每平方米穗数, 是再生稻高产增产的关键。在此基础上, 提出适时早播、畦栽沟灌、二次烤田、重施促芽肥、适高留桩的人工收割高留桩再生稻栽培技术, 并从品种选择、再生季施肥、留桩高度等方面探讨了机械化收割低留桩蓄留再生稻的关键栽培技术及生理生态机制。最后提出了我国再生稻发展亟待解决的几个问题, 认为当前轻简化的机收低留桩再生稻是我国再生稻发展的方向, 并对机收低留桩再生稻的进一步研究作了展望。

... 3 讨论再生季稻产量与头季收割后稻桩上休眠腋芽萌发成穗数密切相关^[16] ...

... 稻根系由两部分组成, 一是母茎上存活的老根与其在再生季产生的新根, 二是稻桩母茎茎节休眠芽上根原基萌发而成的新根^[16], 其中再生新根对机械化收割中低留桩蓄留再生稻再生分蘖的萌发尤为重要, 两者间存在着#cod#x0201c ...

2001

0.0

林文, 李义珍, 姜照伟, 郑景生. 再生稻根系形态和机能的品种间差异及与产量的关联性. 福建农业学报, 2001, (1): 1-4

Lin

, Li Y

, Jiang Z

, Zheng J

. Morphologic and functional difference of root systems among ratooning rice varieties and its correlation with yield. Fujian J Agric Sci, 2001, (1): 1-4 (in Chinese with English abstract)

观察比较了再生稻根系形态和机能的品种间差异以及与产量的关联性，结果表明，汕优明86的根系形态发达，机能高而持久，头季和再生季产量都最高；汕优63的根量和机能次之，双季产量居次；特优70头季根系发达，但成熟期机能骤降，再生季齐穗后根系机能逐渐衰竭，头季高产而再生季低产。说明选用根系形态发达、机能高而持久的品种，是确保再生稻双季稳定高产的基础。 Abstract： The morphologic and functional diference of root systems amongthree ratooning rice varieties and its correlation with yield were studied.The results showed that,Shanyouming 86 had the most flourishing root systems and the highest,lasting function,and its yield in the first season and ratooning season was the highest.In Shanyou 63,its root quantity,function and the yield in both seasons were the second.Teyou 70 had flourishing root system in the first season,but its function sharply dropped after mature period，and gradually went bad after heading stage of the ratooning season so that its yield was high in the first season and low in ratooning season.Therefore,the rice variety with flourishing root systems and high and lasting function will contribute to high and stable yield for ratooning rice.

... 大量的研究结果表明再生稻休眠芽尤其是低节位腋芽的萌发生长成穗与再生季水稻根系活性有着显著或极显著的相关性^[6,17] ...

2012

0.0

... 这与俞道标等的研究结果一致^[18], 也表明了再生季水、肥运筹通过调控再生季水稻的根系活性进而影响再生腋芽的萌发和再生季稻产量 ...

2008

0.0

... 环境条件是通过何种途径调控作物的根系构型呢?张福锁等^[19]认为栽培调控的实质是调控根际环境, 为作物创造一个良好的根际生态环境从而调控作物-土壤-微生物间的相互作用达到促进作物生长和发育的目的, 根际是作物-土壤-微生物及环境条件相互作用的区域, 也是各种营养物质、水分、有益和有害物质及生物作用于根系或进入根系参与循环的界面 ...

2010

0.0

沈建凯, 黄璜, 傅志强, 张亚, 陈灿, 金小马, 廖晓兰, 龙攀, 高文娟, 谢小婷. 规模化稻鸭生态种养对稻田杂草群落组成及物种多样性的影响. 中国生态农业学报, 2010, 18(1): 123-128

Shen J

, Huang

, Fu Z

, Zhang

, Chen

, Jin X

, Liao X

, Long

, Gao W

, Xie X

. Effect of large-scale rice-duck eco-farming on the composition and diversity of weed community in paddy fields. Chin J Eco-Agric, 2010, 18(1): 123-128 (in Chinese with English abstract)

Changing characteristics of weed community, weed-controlling effect of large-scale rice-duck eco-farming (SF), conventional rice-duck eco-farming (CF) compared to mono-cropping rice (CK) were analyzed in order to explore the dynamics of weed communities in large-scale rice-duck eco-farming paddy fields. The study shows that weed density in paddy fields significantly decreases with more than 94% grass-controlling effect under SF compared with CK. Before rice booting stage, weeds compose mainly of Alopecurus pratensis , Echinochhloa crausgalli , Cynodon dactylon , Alternanthera philoxeroides , Mouochoria vaginalis , Murdannia triquetra and Eclipta prostrata in SF. During heading to ripe stage, weeds are mainly A. philoxeroides , E. crausgalli , C. dactylon , M. triquetra . At tillering, species richness, Simpon index and Shannon-Wiener index are higher while Pielou index is lower in SF than in CK. From booting to ripening, weed species richness, Simpon index and Shannon-Wiener index are lower while Pielou index is significantly higher in SF than in CK. In the entire rice growth period, weed density, weed fresh weight and species diversity are not significantly different between SF and CF, but species diversity is higher in SF than in CF. SF changes the structure of weed community and decreases weed species diversity, effectively limiting weed infestation in paddy fields.

为探索规模稻鸭生态种养稻田杂草群落的变化动态, 运用植物群落生态的方法研究了规模稻鸭生态种养、常规稻鸭生态种养和水稻单一种植稻田杂草的群落变化特点及控草效果。结果表明: 规模稻鸭生态种养杂草密度显著降低, 与水稻单一种植比较防效为94%; 水稻分蘖到孕穗期稻田杂草主要由看麦娘( Alopecurus pratensis )、稗草( Echinochhloa crausgalli )、狗牙根( Cynodon dactylon )、水花生( Alternanthera philoxeroides )、鸭舌草( Mouochoria vaginalis )、水竹叶( Murdannia triquetra )和鳢肠( Eclipta prostrata )组成, 抽穗期到成熟期主要由水花生、稗草、狗牙根和水竹叶组成。研究还表明, 规模稻鸭生态种养水稻分蘖期物种丰富度、Simpon指数、Shannon-Wiener指数略高于水稻单一种植, Pielou指数低于水稻单一种植; 孕穗期到成熟期物种丰富度、Simpon指数、Shannon-Wiener指数低于水稻单一种植, Pielou指数显著高于水稻单一种植。水稻全生育期规模稻鸭生态种养与常规稻鸭生态种养比较稻田杂草密度、杂草生物量和多样性指数差异不显著, 但杂草生物多样性指数略有提高。说明规模稻鸭生态种养显著改变稻田杂草的群落结构和组成, 可抑制杂草发生危害, 达到有效控草目的。

... 作物-土壤-微生物及其环境之间的相互作用构成了作物的根际生态系统, 它对于土壤的理化性质、根系的形态、生理有着重要的影响^[20,21], 根际化学过程包括根际土壤的酸碱反应、氧化还原反应等, 根际土壤酶和根际微生物是根际生态系统中重要组成部分, 在根际生态中起着重要的作用, 它们对于植物根际生态系统的物质循环和能量流动具有重要的影响, 由于根际土壤酶和根际微生物对外界条件尤其是土地管理、耕作制度的影响异常敏感, 其变化和群落组成、多样性常常被用作表征土壤质量变化的指标^[22] ...

2013

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裴鹏刚, 张均华, 朱练峰, 禹盛苗, 金千瑜. 根际氧浓度调控水稻根系形态和生理特性研究进展. 中国稻米, 2013, 19(2): 6-8

Pei P

, Zhang J

, Zhu L

, Yu S

, Jin Q

. Research progress on effects of rhizosphere dissolved oxygen on root morphology and physiological characteristics in rice. China Rice, 2013, 19(2): 6-8 (in Chinese)

促进水稻根系生长并保持旺盛的根系活力是提高水稻产量的重要途径。提高根际氧含量，为水稻根系提供良好生长环境，已成为目前国内外根系研究乃至水稻产量形成的热点课题。探明根际氧浓度调控水稻根系生长发育的机理，对于水稻理想根型和高产栽培研究具有重要意义。本文综述了国内外有关根际氧浓度调控水稻根系形态和生理特性的研究进展，提出了在该方向上有待进一步研究的问题与发展趋势，以期为水稻高产栽培的根系调控研究提供依据。

2014

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胡婵娟, 郭雷, 刘国华. 黄土丘陵沟壑区不同植被恢复格局下土壤微生物群落结构. 生态学报, 2014, 34: 2986-2995

Hu C

, Guo

, Liu G

. Soil microbial community structure under different vegetation restoration patterns in the loess hilly area. Acta Ecol Sin, 2014, 34: 2986-2995 (in Chinese with English abstract)

Soil erosion is still severe in the Loess Plateau, and vegetation restoration practices are effective measures to fight it by reducing nutrient loss and improving soil quality. Soil microorganisms are important for soil ecosystem, which play crucial roles in biogeochemical cycling and ecosystem functioning. Soil biological and biochemical properties, such as microbial biomass, microbial community composition, and microbial functional diversity are frequently used for detecting accurate changes in soil properties. The major purpose of this study therefore, is to discover the effects of different vegetation restoration patterns on soil microbial community structures in the Loess Plateau. The study area was located in Yangjuangou catchment, a typical loess hilly area near Yan'an city of northern Shaanxi province, China. Four typical hillslopes, including F (artificial forest), G (grass), G-F-G (grass on the upper and lower slopes and artificial forest on the middle slope), and F-G-F (artificial forest on the upper and lower slopes and grass on the middle slope), were chosen. Several soil sampling sites, each covered by an area of 200m 2 , were conducted on the summit, shoulder, backslope, footslope and toeslope along each hillslope, respectively. Three experimental plots in each site were conducted with an area of 25 m 2 . Soil samples in the first two vertical layers (0-10 cm and 10-20 cm) were collected in August of 2007, for analyzing soil microbial community structure and soil physicochemical properties. Soil microbial community structure was analyzed by using phospholipid fatty acid (PLFA) analytical method and soil physicochemical properties were done by general soil and agricultural chemistry analytical methods. Results showed that significant difference of phospholipid fatty acid composition existed between different soil layers and soils with different vegetation restoration patterns, mainly in surface soil layer collected from G-F-G and F-G-F differing with that from F and G. The proportion of phospholipid fatty acid indicated fungi was higher in the surface soils under G-F-G and F-G-F, compared to F and G. Principal components analysis (PCA) showed that in the 0-10 cm soil layer, the phospholipid fatty acid structure of G-F-G was different with that of F and G, while the phospholipid fatty acid structure of F-G-F was different with F significantly. The biomass of soil bacteria, fungi, Gram-negative bacteria and Gram-positive bacteria under F-G-F pattern were all captured the highest values. There was significant difference of soil bacteria biomass and Gram-positive bacteria biomass in the surface soil layers among the four mentioned vegetation restoration patterns. The biomass of soil bacteria and fungi, however, had non-significant difference among 10-20 cm soil layers with different vegetation restoration patterns. Redundancy analysis (RDA) showed that soil organic carbon and total nitrogen had more important relationship with soil microbial community. However, the effects of some physicochemical properties such like pH and bulk density on soil microbial community needed more study. In conclusion, microbial community structure was dissimilar in soils and significant difference mainly appeared in surface soil layers under four vegetation restoration patterns. The roles of artificial forestation and forest-grass collocation on the slope in improving soil microbial biomass were believed much better than abandoned farmland.

针对典型黄土丘陵沟壑区陕西延安羊圈沟小流域坡面上单一刺槐林、单一撂荒草地以及林草搭配的草地-林地-草地及林地-草地-林地4种不同植被格局，利用磷脂脂肪酸（phospholipid fatty acid，PLFA）谱图分析法对土壤微生物群落结构进行监测研究，旨在揭示坡面上不同的植被恢复格局对土壤微生物群落结构的影响。研究发现4种不同植被格局下，2种林草搭配的植被格局磷脂脂肪酸的结构比较相似，与单一植被格局相比，表层土壤中表征真菌的特征脂肪酸所占的比例有所提高。主成分分析显示4种植被格局0-10 cm土壤微生物群落结构存在差异，差异主要存在于2种林草搭配的植被格局与2种单一的植被格局之间，其中草地-林地-草地的植被格局与刺槐林和撂荒草地之间土壤微生物群落结构的差异均达到了显著水平。不同微生物菌群的量在4种植被格局土壤间显著性差异主要存在于表层土壤中的细菌菌群和革兰氏阳性菌，革兰氏阴性菌和真菌在4种植被格局土壤之间无显著差异。总之，4种不同植被恢复格局的土壤微生物群落结构存在差异且差异主要存在于表层土壤，坡面上人工林的种植及林草搭配的恢复模式较直接撂荒更有利于提高微生物菌群的生物量。

2004

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... Waldrop等^[23]的研究结果也指出土壤多酚氧化酶和过氧化物酶的活性对于土壤水分的变化比较敏感, 水分的增加会降低这两种酶的活性, 与G处理相比, GN处理对于六大土壤酶活性都有显著提高作用, 这与周卫等^[24]和赵之重^[25]的报道相似, 表明再生季干湿交替灌溉有利于提高土壤的通透性和氧含量, 对于根际土壤中有机质的氧化和腐殖质的形成有促进作用, 而水肥耦合措施对于根际土壤有机质的氧化和腐殖质的形成及增加根际土壤中养分有效性的促进作用要显著高于单一措施, 从而为再生季新根的形成和腋芽的萌发提供相应的物质需求保障 ...

1990

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1998

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1998

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高子勤, 张淑香. 连作障碍与根际微生态研究: I. 根系分泌物及其生态效应. 应用生态学报, 1998, 9: 549-554

Gao Z

, Zhang S

. Continuous cropping obstacle and rhizospheric microecology: I. Root exudates and their ecological effects. Chin J Appl Ecol, 1998, 9: 549-554 (in Chinese with English abstract)

Growth degeneration and yield declination are becoming obvious after a long term continuous cropping of crops, vegetables, fruit trees and nursery seedlings.Many studies show that soil ecological environment greatly affect the growth of plant under continuous cropping, and particularly, the decomposition products of plant residues and pathogenic microbes have a toxic effect to plants and affect the normal metabolism of root exudates, producing an auto-toxic effect.In this paper, the rhizospheric effects of soil environmental factors (air, moisture, nutrient and microorganisms), active substances (phytotoxin, residue decomposition substances, microbial toxin), and pathogenic microbes on root exudates under conditions of continuous cropping were systematically introduced, which would afford inspiration for further studying on the interaction mechanism between root exudates and continuous cropping obstacle.

Institute of Applied Ecology, Academia Sinica, Shenyang 110015

作物、蔬菜、果树以及苗木长期连作后,皆出现生长衰退和产量降低。许多研究结果表明,连作条件下土壤生态环境对植物生长有很大的影响,尤以植物残体与病原微生物的分解产物,对植物有致毒作用,并影响植物根系分泌物正常代谢,以致于发生自毒作用。本文围绕根系分泌物与根际微生态的相互关系,系统地介绍连作障碍条件下,影响根系分泌物的环境因素(土壤空气、湿度、养分与微生物)、活性物质(自身毒素、残体分解物、微生物产生毒素)、土壤病原菌等的根际效应,为深入研究根系分泌物与连作障碍的相互作用机制提供启示。

... 1%, 在两大成分分离中起主要贡献作用的是聚合物类、碳水化合物类和氨基酸类, 土壤碳水化合物类和氨基酸类是土壤中微生物细胞必需的组成物质和活动的主要能源, 不同水肥调控下根际土壤微生物对根系环境中碳水化合物和氨基酸的利用程度不同, 直接影响根际土壤微生物的数量和种类, 根际土壤微生物数量和种类的不同变化又反过来影响着作物根系的生长发育^[26,27] ...

2000

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