Effects of localized nitrogen supply treatments on growth and root parameters in Pinus massoniana families under phosphorus deficiency
SONGPing1, ZHANGRui1,*,, ZHOUZhi-Chun1, TONGJian-She2, WANGHui2 1Research Institute of Subtropical Forestry, Chinese Academy of ForestryState Forestry Administration Engineering Research Center of Masson PineZhejiang Provincial Key Laboratory of Tree Breeding, Hangzhou 311400, Chinaand 2Fuxi Forest Farm, Chun’an, Zhejiang 311701, China; 通讯作者:* 通信作者Author for correspondence (E-mail: ruirui0218@126.com) 收稿日期:2017-01-3 接受日期:2016-08-9 网络出版日期:2017-07-28 版权声明:2017植物生态学报编辑部本文是遵循CCAL协议的开放存取期刊，引用请务必标明出处。 基金资助:基金项目 “十二五”国家科技支撑计划项目(2012BAD01B02)、国家自然科学基金(31370671)和浙江省农业(林木)新品种选育重大专项课题(2016C02056-4)
关键词：家系;局部供氮;低磷胁迫;马尾松;根系参数 Abstract Aims A heterogeneous spatially distribution of nutrients in natural soil may affect plant growth. The objective of this study was to determine the effects of localized nitrogen (N) supply treatments on growth traits and root parameters among different families in Pinus massoniana.Methods Five families of P. massoniana seedlings from full-sib progenies were used as test materials (1, 25, 49, 52, and 57). This study included two conditions, (i.e. homogeneous phosphorus (P) deficiency vs. heterogeneous P efficiency) among soil layers in combination with four N supply treatments in a one-year pot experiment. These N supply treatments were: (1) Homogeneously high N along the soil profile (HHH); (2) high N-high N-low N (HHL); (3) low N-low N-high N (LLH); (4) low N-low N-on side with N addition and the other side without N supply (LLH/L).Important findings This study indicated that localized N supply treatment did enhance the growth of P. massoniana, and this enhancement mainly happened in the pattern of N applied to deep soil. The results showed: 1) Compared to the homogeneous low P condition, there were increase in the growth traits and root parameters of P. massoniana under heterogeneous low P condition. Particularly, the root length and root surface area under the heterogeneous P deficiency condition were 1.95 times and 2.11 times higher than that subjected to the homogeneous P deficiency. 2) Localized N supply treatment affected seedling growth, and there was a significant interaction among N supply pattern and P condition. In compared with homogeneous N supply treatment, the height, basal diameter and dry weight of seedlings increased significantly by localized N supply treatments (LLH and/or LLH/L) under both two P deficiency conditions. But when the seedlings parameters were enhanced under homogeneous P deficiency, they were inhibited under heterogeneous P deficiency subjected to HHL. 3) Within the two P conditions, LLH and LLH/L stimulated root proliferation significantly, and root parameters were significantly enhanced under the heterogeneous P deficiency condition. Specifically, the root length and root surface area subjected to LLH/L rather than HHH were significantly enhanced by 29.2% and 32.3%, respectively. However, the length and surface area of the roots were suppressed by HHL treatment. 4). There were significant differences in response to different N supply treatments among P. massoniana families Seedlings in the families of 49, 52, and 57 responded to the localized N supply treatments with increased root proliferation, which enhanced seedling dry mass. On the other hand, the seedling growth in the family of 25 were stimulated by N and (or) P concentration, while the response of seedlings in the family of 1 to local nitrogen supply was relatively slow and exhibited growth retardation.
Keywords：family;localized nitrogen supply;phosphorus deficiency;Pinus massoniana; root parameter -->0 PDF (1195KB)元数据多维度评价相关文章收藏文章 本文引用格式导出EndNoteRisBibtex收藏本文--> 宋平, 张蕊, 周志春, 童建设, 王晖. 局部供氮对低磷胁迫下马尾松不同家系生长及根系参数的影响. 植物生态学报, 2017, 41(6): 622-631 https://doi.org/10.17521/cjpe.2016.0260 SONGPing, ZHANGRui, ZHOUZhi-Chun, TONGJian-She, WANGHui. Effects of localized nitrogen supply treatments on growth and root parameters in Pinus massoniana families under phosphorus deficiency. Chinese Journal of Plant Ecology, 2017, 41(6): 622-631 https://doi.org/10.17521/cjpe.2016.0260 土壤养分在空间分布上多呈时空异质性或者斑块分布(Hutching & John, 2004)。产生土壤养分空间异质性的原因很多, 其中包括有机物(凋落物、根系死亡以及动物尸体)的分解、无机养分在土壤中的扩散系数差异以及微地形因素影响等(Hodge, 2004; 徐星凯等, 2012; 刘双娥等, 2015)。植物为了获得更多的养分, 其根系会增加在土壤养分富集区域的分布比例, 进而提高根系对养分的获取和利用能力(Hodge, 2004; Mommer et al., 2012)。近年来, 随着合理高效的供肥管理措施(局部施肥)在农业中广泛应用及其高效的生产力, 使其在林业上尤其是人工林经营管理中也开始得到广泛应用。因此, 了解肥料集约经营管理下的林木根系生长、形态分布以及养分吸收状况, 将有利于解释林木在生长上存在的差异, 同时为提高速生丰产林生产力提供更为完善和高效的技术指导。 氮和磷是植物生长最重要的两种营养元素, 其有效性和空间分布对植物的生长和根系发育具有重要影响(Hodge, 2004; Vance et al., 2003)。研究揭示, 局部供氮或供磷可显著促进大麦(Hordeum vulgare)、玉米(Zea mays)和小麦(Triticum aestivum)等植物侧根数量的增加以及总根长的提高(Drew, 1975; Officer et al., 2009)。Zhang和Forde (2000)将拟南芥(Arabidopsis thaliana)分别放在局部高氮和均匀高氮的琼脂培养基上培养时发现, 根系会在富氮斑块中增生, 而均匀供氮处理中的根系总量降低。可见根系在富养斑块中的增生可促进植物对养分供应的有效利用(Linkohr et al., 2002)。更重要的是, 相比于单独局部供氮或供磷, 在斑块中同时添加氮和磷更能促进根系在富养斑块中增生, 提高营养元素的吸收效率, 进而加快整株的生长(Jing et al., 2010)。这表明植物根系在局部供养斑块中的增生受到营养元素组成以及元素之间互作效应的共同影响。在木本植物研究中也发现类似的变化规律, 但物种间差异较大(Hodge, 2004)。Einsmann等(1999)发现, 10种木本植物中有4种植物的生物量在养分异质性的供应条件下有显著提高, 但在对同种植物的不同研究中, Bliss等(2002)并没有发现这一趋势。虽然有研究认为养分斑块的存在能够对植物地上部分的生长产生积极的促进效应(Jackson & Caldwell, 1996), 但多年生植物绒毛草(Holcus lanatus)仅在短期内出现大量根系增生和促进地上部分生长的现象, 长期来看这种优势很快会消失, 甘松茅(Nardus stricta)则对养分斑块反应不敏感(Fransen & de Kroon, 2001)。显然, 养分斑块或者局部养分供应对植物生长的影响受很多因素的影响, 除了养分的属性外, 植物对养分的敏感性及试验中植物生长时间的长短对结果都会产生较大影响(李洪波等, 2013)。 马尾松(Pinus massoniana)是重要的材用、脂用和生态造林树种, 广泛分布于我国的亚热带地区(秦国峰和周志春, 2012)。当前研究已揭示了氮添加对低磷胁迫下马尾松生长影响的生物学机制, 包括地上部分生长、根系参数和氮、磷效率的适应性变化等(Zhang et al., 2013; 庞丽等, 2014)。 然而, 相关研究未考虑氮素本身在空间分布上的不均一性。鉴于此, 本文拟通过人工控制氮素在土壤空间上的异质性, 研究低磷胁迫下马尾松不同家系在局部供氮条件下的生长表现和根系发育, 进一步阐明马尾松不同家系对土壤异质养分的响应差异及机制, 为马尾松营养高效新品种选育和苗木高效栽培方式提供理论依据以及实践指导。
1 材料和方法
1.1 供试材料
试验在浙江省淳安县富溪林场温室大棚内进行(119.13° E, 29.48° N), 其所处海拔为130 m, 年平均气温17.5 ℃, 年降水量为1β701.2 mm。以马尾松二代育种群体内控制授粉产生的5个全同胞子代1、25、49、52和57作为试验材料。盆栽容器采用定制的圆柱形无纺布植树袋, 直径为12 cm, 高28 cm。土壤基质为浙江省淳安县千岛湖的贫瘠酸性红壤, 其理化性质见表1。 Table 1 表1 表1盆栽土壤基质理化性质 Table 1Soil physical and chemical properties of the substrate in the pot experiment
1.2.1 试验装置 本试验中的马尾松分层隔网盆栽设计, 参考杨青等(2011)的磷控释土培系统, 同时结合一年生马尾松苗木的根系形态特征进行分层处理(图1)。盆栽容器自上而下依次分为表层、中间层和深层3个层次, 间距8 cm, 每层之间用孔径为3 mm的塑料纤维网隔离(主根、基根均能顺利穿过)。在试验装置上端填充4 cm厚的干净河沙作为缓冲层, 将经过0.2%- 0.4%的高锰酸钾溶液浸泡20 min的马尾松不同家系种子植入缓冲层中间, 每盆播种在3-5粒种子。 显示原图|下载原图ZIP|生成PPT 图1不同磷环境下的局部供氮处理。HHH, 均匀供氮; HHL, 表层和中间层供氮; LLH, 底层供氮; LLH/L, 底层半侧供氮。 -->Fig. 1The patterns of N added under two P deficiency conditions. HHH, homogeneously high N along the soil profile; HHL, high N-high N-low N; LLH, low N-low N-high N; LLH/L, low N-low N-on side with N addition and the other side without N supply. -->
方差分析结果表明, 在两种低磷环境下, 局部供氮对马尾松苗木的株高、地径和干物质积累量均有显著影响(p < 0.001)(表2)。与同质低磷相比, 异质低磷下的株高、地径和干物质积累量分别增加了5.1%、11.6%和22.0%, 其中干物质积累量之间的差异达到显著水平(p < 0.05), 但在HHL处理下的生长表现例外(表2)。与均匀供氮(HHH)相比, HHL处理显著增加了同质低磷下马尾松苗木的株高、地径和干物质积累量, 相反, HHL处理却不利于异质低磷下马尾松苗木的生长, 其干物质积累量分别在同质低磷下增加了41.7%, 异质低磷下降了36.9% (表2)。LLH和LLH/L处理后, 马尾松苗木的株高、地径和干物质积累量与HHH处理后的生长表现无显著差异, 说明在底层或底层半侧较少氮素投入的情况下, 马尾松苗木可获得与均匀供氮同样的生长表现。另外, 同质低磷下的根冠比小于异质低磷水平, 且在局部供氮处理后进一步降低, 表明局部供氮对同质低磷下马尾松苗木地上部分的生长更为有利(表2)。 Table 2 表2 表2局部供氮对马尾松生长、根系参数和氮磷含量的影响及方差分析(平均值±标准误差) Table 2Effects of localized N supply treatments on growth traits, root parameters and N and P concentration of Pinus massoniana (mean ± SE)
磷环境 P level
性状 Trait
氮处理 N supply treatment
变异来源 Source of variation (F值 F value)
HHH
HHL
LLH
LLH/L
家系 Family
氮处理 N
家系&×氮处理 Family &× N
同质低磷 Homo low P
SH (cm)
14.42 &± 0.52b
16.92 &± 0.54a
14.90 &± 0.65b
14.66 &± 0.65b
47.73***
6.34***
2.13*
SBD (mm)
2.39 &± 0.10b
2.99 &± 0.07a
2.61 &± 0.09b
2.61 &± 0.09b
23.96***
12.50***
2.24*
SDW (g)
1.32 &± 0.12b
1.87 &± 0.10a
1.44 &± 0.13b
1.63 &± 0.12ab
19.21***
6.11**
2.74*
RDW (g)
0.26 &± 0.02b
0.37 &± 0.03a
0.27 &± 0.02b
0.35 &± 0.02a
12.02***
7.49***
1.54
R/S
0.36 &± 0.03a
0.17 &± 0.03b
0.23 &± 0.03ab
0.18 &± 0.03b
12.23***
2.75*
2.55*
RL (cm)
192.36 &± 9.72a
186.38 &± 11.93a
210.54 &± 14.45a
199.73 &± 11.92a
12.28***
2.92*
1.20
RSA (cm2)
47.37 &± 3.00b
46.50 &± 2.56b
61.96 &± 4.38a
59.90 &± 3.65a
13.14***
7.02***
1.47
PC (mg&·g-1)
0.75 &± 0.03b
1.14 &± 0.04a
0.87 &± 0.08ab
0.95 &± 0.11ab
1.02
0.96
1.02
NC (mg&·g-1)
17.02 &± 0.39b
21.68 &± 2.64a
18.89 &± 0.34ab
16.18 &± 0.59b
3.44**
3.37*
1.47
异质低磷 Hetero low P
SH (cm)
17.51 &± 0.74a
13.62 &± 0.52b
16.72 &± 0.67a
16.13 &± 0.67a
43.10***
12.58***
2.54*
SBD (mm)
3.18 &± 0.11a
2.61 &± 0.11c
3.11 &± 0.10ab
2.82 &± 0.12bc
17.46***
8.22***
2.06*
SDW (g)
2.25 &± 0.19a
1.42 &± 0.13b
2.01 &± 0.14a
1.96 &± 0.16a
15.53***
6.34***
2.32*
RDW (g)
0.47 &± 0.04a
0.36 &± 0.03a
0.42 &± 0.04a
0.47 &± 0.06a
13.33***
1.97
1.04
R/S
0.26 &± 0.03a
0.34 &± 0.05a
0.26 &± 0.02a
0.31 &± 0.04a
7.30***
0.65
0.49
RL (cm)
368.77 &± 27.72ab
296.81 &± 21.57b
399.17 &± 31.82ab
476.60 &± 61.51a
6.70***
4.13**
1.09
RSA (cm2)
105.77 &± 7.81b
94.15 &± 7.77b
115.78 &± 9.16ab
139.34 &± 18.01a
8.22***
3.23*
1.04
PC (mg&·g-1)
0.90 &± 0.03a
1.18 &± 0.04a
1.02 &± 0.03a
1.11 &± 0.08a
1.05
1.06
1.04
NC (mg&·g-1)
17.33 &± 0.39b
18.46 &± 0.34a
15.23 &± 0.41c
15.17 &± 0.39c
15.77***
23.80***
1.36
SH, 苗高; SBD, 地径; SDW, 干物质量; RDW, 根干物质量; R/S, 根冠比; RL, 根长; RSA, 根表面积; PC, 全磷含量; NC, 全氮含量。 *, 0.05 > p > 0.01; **, 0.01 > p > 0.0001; ***, p < 0.0001。 SH, seedling height; SBD, seedling basal diameter; SDW, seedling dry mass; RDW, root dry mass; R/S, root/shoot ratio; RL, root length; RSA, root surface area; PC, phosphorus concentration; NC, nitrogen concentration. *, 0.05 > p > 0.01; **, 0.01 > p > 0.000β1; ***, p < 0.000 1. 新窗口打开 不同家系对局部供氮的响应差异显著(p < 0.05) (表2)。如图2所示, 在两种低磷环境下, 家系1在各氮处理后的株高和干物质积累量均显著低于其他供试家系。家系25在同质低磷、HHL处理后, 其株高和干物质积累量较HHH处理显著高出23.3%和57.0%; 但在异质低磷下, LLH和LLH/L处理更有利于该家系的生长。家系49和52的干物质积累量在同质低磷、LLH处理下较HHH处理分别降低了33.9%和15.6%, 但在异质低磷、LLH处理下与HHH处理之间无显著差异。家系57的株高和干物质积累量的变化与家系49以及52的变化规律类似, 且较其他家系具有更强的生长势, 尤其在同质低磷环境下更为突出。 显示原图|下载原图ZIP|生成PPT 图2局部供氮对低磷胁迫下马尾松不同家系苗木生长的影响(平均值±标准误差)。HHH, 均匀供氮; HHL, 表层和中间层供氮; LLH, 底层供氮; LLH/L, 底层半侧供氮。不同小写字母表示不同处理之间差异显著(p < 0.05)。 -->Fig. 2The seedling growth traits of Pinus massoniana families in different local N supply treatments (mean ± SE). HHH, homogeneously high N along the soil profile; HHL, high N-high N-low N; LLH, low N-low N-high N; LLH/L, low N-low N-on side with N addition and the other side without N supply. Different lower letters indicate significant differences among treatments (p < 0.05). -->
2.2 局部供氮对不同磷环境下马尾松不同家系苗木根系参数的影响
马尾松苗木的根系参数对局部供氮的响应在两种低磷环境下类似, 底层供氮皆有利于苗木根系的生长。与HHH相比, LLH和LLH/L处理促进了马尾松苗木根长和根表面积的增加, 尤其在异质低磷、LLH/L处理后, 其根长和根表面积显著增加了29.2%和32.3%; 相反, HHL处理降低了马尾松苗木的根长和根表面积(表2)。异质低磷下的根长和根表面积约是同质低磷下的1.95倍和2.11倍, 说明异质低磷较同质低磷显著促进了根系的生长发育(p < 0.01) (表2)。 不同家系的根系参数对局部供氮的响应程度有所差异。由图3可知, 在两种低磷环境下, 家系25的根长和根表面积在不同供氮处理后均无明显变化, 但在同质低磷、HHL处理后, 该家系的根长和根表面积分别降低了13.4%和25.8% (p = 0.07)。家系49、52和57在同质低磷下, 经LLH和LLH/L处理后, 其根长和根表面积较HHH处理有所下降, 但未达到显著水平; 相反, 在异质低磷下, LLH和LLH/L处理后的根长和根表面积明显高于HHH处理水平, HHL处理则不利于根系的生长发育。家系1的根长和根表面积的变化与生长性状的变化规律相似, 均显著小于其他家系。 显示原图|下载原图ZIP|生成PPT 图3局部供氮对低磷胁迫下马尾松不同家系苗木根系参数的影响(平均值±标准误差)。HHH, 均匀供氮; HHL, 表层和中间层供氮; LLH, 底层供氮; LLH/L, 底层半侧供氮。不同小写字母表示不同处理之间差异显著(p < 0.05)。 -->Fig. 3The root parameters of Pinus massoniana families in different local N supply treatments (mean ± SE). HHH, homogeneously high N along the soil profile; HHL, high N-high N-low N; LLH, low N-low N-high N; LLH/L, low N-low N-on side with N addition and the other side without N supply. Different lower letters indicate significant differences among treatments (p < 0.05). -->
2.3 局部供氮对不同磷环境下马尾松不同家系苗木氮、磷含量的影响
马尾松苗木植株的全氮含量受局部供氮处理影响显著, 全磷含量无显著差异, 但在异质低磷下的全磷含量显著高于同质低磷水平(表2)。如图4显示, 在异质低磷下, 马尾松苗木的全磷含量平均高于同质低磷水平13.5%, 全氮含量却平均下降了10.3%。与HHH处理相比, 在两种低磷环境下, 局部供氮(HHL、LLH和LLH/L)处理后, 马尾松苗木植株的全磷含量均有不同程度的增加, 全氮含量则因不同的供氮方式差异较大, 其中, HHL处理显著提高了马尾松苗木植株的全氮含量 (同质低磷下较HHH增加了27.4%; 异质低磷下增加了6.5%), LLH和LLH/L处理则不利于苗木植株全氮含量的增加, 甚至显著降低了异质低磷下苗木的全氮含量。 显示原图|下载原图ZIP|生成PPT 图4局部供氮对两种磷环境下马尾松不同家系氮、磷含量的影响(平均值±标准误差)。HHH, 均匀供氮; HHL, 表层和中间层供氮; LLH, 底层供氮; LLH/L, 底层半侧供氮。不同小写字母表示不同处理之间差异显著(p < 0.05)。不同小写字母表示不同处理之间差异显著(p < 0.05)。 -->Fig. 4N concentration and P concentration of P. massoniana families in different local N supply treatments (mean ± SE). HHH, homogeneously high N along the soil profile; HHL, high N-high N-low N; LLH, low N-low N-high N; LLH/L, low N-low N-on side with N addition and the other side without N supply. Different lower letters indicate significant differences among treatments (p < 0.05). -->
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Are competitive interactions influenced by spatial nutrient heterogeneity and root foraging behavior? 2002
Comparison of effects of a localized supply of phosphate, nitrate, ammonium and potassium on growth of seminal root system and shoot in barley. 1 1975
... 局部供氮对植物根系形态和分布的影响较大(Li et al., 2014).研究结果表明, 在两种低磷环境下, 与HHH相比, LLH/L对马尾松苗木根系的生长发育有明显的促进作用, 尤其在异质低磷土壤环境下的促进作用更为突出, 其次是LLH, HHL反而不利于根系的生长发育.分析认为, 底层半侧供氮有利于根系增生发育的原因可能在于: 植物根系感知一侧氮信号的缺乏, 诱导了根原基和初级根的形成, 而富氮斑块中充足的氮供应则加速了根系的伸长(He et al., 2003; Li et al., 2014), 进而促进了植物整体根长和根表面积的增加.如在异质低磷下, LLH/L处理中的总根长是LLH处理的1.30倍.相似的规律在水稻感知局部磷胁迫的研究中也被发现(He et al., 2003).另一方面, HHL对根长和根表面积有一定的抑制效应, 推测这可能是由于: 首先, 马尾松本身属于深根型植物, 根的向地性促使其向土壤深处觅取更多的养分, 而HHL处理中的底层并没有充足的氮素供应, 限制了根系的生长发育.这一结果与前人对禾本科植物的研究结果有所差异, 如小麦、玉米和水稻(Oryza sativa)等植物可通过增加根系在土壤表层富养区域的分布来响应自然界土壤中的异质养分环境(Weligama et al., 2008; Jing et al., 2010; Li et al., 2014).马尾松作为木本植物, 其对土壤养分异质性分布的响应可能因树种和土壤养分组成的不同而差异较大(Einsmann et al., 1999; Guo et al., 2002; 梅莉等, 2006; 杨青等, 2011); 其次, 该实验的种子播种于土壤表层, 其生长初期的根系增生和扩张及时获得了营养的补充, 使得植物在养分分配时, 将更多的营养元素输送到了其他器官(茎、叶), 降低了对根系的养分投入, 限制了根系的增生发育(段洪浪等, 2009; 吴茜等, 2011).结合苗期马尾松的株高、地径等地上部分的生长状况也印证了这一推测. ...
Nutrient forging traits in 10 co-occurring plant species of contrasting life forms. 1 1999
... 局部供氮对植物根系形态和分布的影响较大(Li et al., 2014).研究结果表明, 在两种低磷环境下, 与HHH相比, LLH/L对马尾松苗木根系的生长发育有明显的促进作用, 尤其在异质低磷土壤环境下的促进作用更为突出, 其次是LLH, HHL反而不利于根系的生长发育.分析认为, 底层半侧供氮有利于根系增生发育的原因可能在于: 植物根系感知一侧氮信号的缺乏, 诱导了根原基和初级根的形成, 而富氮斑块中充足的氮供应则加速了根系的伸长(He et al., 2003; Li et al., 2014), 进而促进了植物整体根长和根表面积的增加.如在异质低磷下, LLH/L处理中的总根长是LLH处理的1.30倍.相似的规律在水稻感知局部磷胁迫的研究中也被发现(He et al., 2003).另一方面, HHL对根长和根表面积有一定的抑制效应, 推测这可能是由于: 首先, 马尾松本身属于深根型植物, 根的向地性促使其向土壤深处觅取更多的养分, 而HHL处理中的底层并没有充足的氮素供应, 限制了根系的生长发育.这一结果与前人对禾本科植物的研究结果有所差异, 如小麦、玉米和水稻(Oryza sativa)等植物可通过增加根系在土壤表层富养区域的分布来响应自然界土壤中的异质养分环境(Weligama et al., 2008; Jing et al., 2010; Li et al., 2014).马尾松作为木本植物, 其对土壤养分异质性分布的响应可能因树种和土壤养分组成的不同而差异较大(Einsmann et al., 1999; Guo et al., 2002; 梅莉等, 2006; 杨青等, 2011); 其次, 该实验的种子播种于土壤表层, 其生长初期的根系增生和扩张及时获得了营养的补充, 使得植物在养分分配时, 将更多的营养元素输送到了其他器官(茎、叶), 降低了对根系的养分投入, 限制了根系的增生发育(段洪浪等, 2009; 吴茜等, 2011).结合苗期马尾松的株高、地径等地上部分的生长状况也印证了这一推测. ...
Long-term disadvantages of selective root placement: Root proliferation and shoot biomass of two perennial grass species in a 2-year experiment. 1 2001
Localized application of phosphorus and ammonium improves growth of maize seedlings by stimulating root proliferation and rhizosphere acidification. 3 2010
Nutrient uptake, cluster root formation and exudation of protons and citrate in Lupinus albus as affected by localized supply of phosphorus in a split-root system. 1 2005
... 局部供氮对植物根系形态和分布的影响较大(Li et al., 2014).研究结果表明, 在两种低磷环境下, 与HHH相比, LLH/L对马尾松苗木根系的生长发育有明显的促进作用, 尤其在异质低磷土壤环境下的促进作用更为突出, 其次是LLH, HHL反而不利于根系的生长发育.分析认为, 底层半侧供氮有利于根系增生发育的原因可能在于: 植物根系感知一侧氮信号的缺乏, 诱导了根原基和初级根的形成, 而富氮斑块中充足的氮供应则加速了根系的伸长(He et al., 2003; Li et al., 2014), 进而促进了植物整体根长和根表面积的增加.如在异质低磷下, LLH/L处理中的总根长是LLH处理的1.30倍.相似的规律在水稻感知局部磷胁迫的研究中也被发现(He et al., 2003).另一方面, HHL对根长和根表面积有一定的抑制效应, 推测这可能是由于: 首先, 马尾松本身属于深根型植物, 根的向地性促使其向土壤深处觅取更多的养分, 而HHL处理中的底层并没有充足的氮素供应, 限制了根系的生长发育.这一结果与前人对禾本科植物的研究结果有所差异, 如小麦、玉米和水稻(Oryza sativa)等植物可通过增加根系在土壤表层富养区域的分布来响应自然界土壤中的异质养分环境(Weligama et al., 2008; Jing et al., 2010; Li et al., 2014).马尾松作为木本植物, 其对土壤养分异质性分布的响应可能因树种和土壤养分组成的不同而差异较大(Einsmann et al., 1999; Guo et al., 2002; 梅莉等, 2006; 杨青等, 2011); 其次, 该实验的种子播种于土壤表层, 其生长初期的根系增生和扩张及时获得了营养的补充, 使得植物在养分分配时, 将更多的营养元素输送到了其他器官(茎、叶), 降低了对根系的养分投入, 限制了根系的增生发育(段洪浪等, 2009; 吴茜等, 2011).结合苗期马尾松的株高、地径等地上部分的生长状况也印证了这一推测. ...
Regulation of Arabidopsis root development by nitrate availability. 2000
Genetic variations in root morphology and phosphorus efficiency of Pinus massoniana under heterogeneous and homogeneous low phosphorus conditions. 1 2013
, 周志春
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