植物生态学报 ›› 2021, Vol. 45 ›› Issue (7): 714-727.DOI: 10.17521/cjpe.2021.0160
李孝龙1,4, 周俊1,*(), 彭飞2, 钟宏韬3, Hans LAMBERS3
收稿日期:
2021-04-23
接受日期:
2021-06-15
出版日期:
2021-07-20
发布日期:
2021-10-22
通讯作者:
周俊 ORCID:0000-0001-7315-6645
作者简介:
* 周俊: ORCID: 0000-0001-7315-6645, zhoujun@imde.ac.cn基金资助:
LI Xiao-Long1,4, ZHOU Jun1,*(), PENG Fei2, ZHONG Hong-Tao3, Hans LAMBERS3
Received:
2021-04-23
Accepted:
2021-06-15
Online:
2021-07-20
Published:
2021-10-22
Contact:
ZHOU Jun ORCID:0000-0001-7315-6645
Supported by:
摘要:
自然成土过程中土壤养分的变化与植被原生演替常同时发生。随成土年龄变化的植物养分捕获策略(NASs)对植物竞争能力和演替过程具有重要影响。该文将植物NASs划分为细根、微生物、特殊根系、食虫和寄生策略等5个类型; 发现植物NASs的多样性随成土年龄的增加呈哑铃型变化模式; 特殊根系策略对植物捕获养分的作用在成土中期最小、后期最大, 细根和微生物策略的作用随成土年龄的增加逐渐降低; 分析了成土过程中NASs对植物种间关系影响的变化, 发现NASs对成土早期植物的促进作用和中期的竞争关系具有重要影响, 而成土后期多样和互补的NASs对植被群落的稳定共存及多样性的形成具有影响; 提出应进一步探究成土过程中土壤养分与植物NASs变化之间的定量关系, 开展更多研究以阐明NASs对植被原生演替、物种多样性形成和成土过程的贡献与机理。
李孝龙, 周俊, 彭飞, 钟宏韬, Hans LAMBERS. 植物养分捕获策略随成土年龄的变化及生态学意义. 植物生态学报, 2021, 45(7): 714-727. DOI: 10.17521/cjpe.2021.0160
LI Xiao-Long, ZHOU Jun, PENG Fei, ZHONG Hong-Tao, Hans LAMBERS. Temporal trends of plant nutrient-acquisition strategies with soil age and their ecological significance. Chinese Journal of Plant Ecology, 2021, 45(7): 714-727. DOI: 10.17521/cjpe.2021.0160
图2 几种植物养分捕获策略的形态示意图。A, 丛枝菌根真菌Glomus caledonium的菌丝体从豆科宿主Trifolium repens的根系向土壤生长。B, 微宇宙培育的松科植物Pinus sylvestris的幼苗与外生菌根真菌Suillus bovinus共生。可以看到在微宇宙中真菌菌丝在土壤中扩散(箭头头部)并且局部增殖形成清晰致密的斑块(箭头)。C, Woollsia pungens的欧石南类菌根表皮被欧石南类菌根真菌的菌丝套侵染(箭头表示被蓝色染色)。D, 豆科植物Astragalus mahoshanicus的根瘤。E, 山龙眼科植物Banksia grandis的排根。图片由钟宏韬拍摄。F, 杜鹃花科植物Actinocephalus cabralensis的固沙根。G, 莎草科植物Tetraria的胡萝卜状根。H, Velloziaceae植物Barbacenia tomentosa田间采集的毛状根。图A引自Olsson等(2002); 图B、C、G引自Lambers等(2008); 图F引自Oliveira等(2015); 图H引自Abrahão等(2020)。
Fig. 2 Morphology of plant nutrient-acquisition strategies. A, Hyphae of the arbuscular mycorrhizal fungus Glomus caledonium growing into soil from a host root of Trifolium repens (Fabaceae). Photo by Iver Jakobsen; reprinted with permission of Springer-Verlag (Olsson et al., 2002). B, A seedling of Pinus sylvestris (Pinaceae) growing in a microcosm in association with the ectomycorrhizal fungus Suillus bovinus. The fungal mycelium can be seen spreading in the soil in the microcosm (arrowhead) and proliferating locally to form well-defined dense patches (arrows). Reprinted with permission of Elsevier B.V. (Lambers et al., 2008). C, Ericoid mycorrhizal root of Woollsia pungens, showing epidermal cells colonized by coils of an ericoid mycorrhizal fungus (stained blue, arrowed). Reprinted with permission of Elsevier B.V. (Lambers et al., 2008). D, Nodules of Astragalus mahoshanicus (Fabaceae). E, Cluster roots of Banksia grandis (Proteaceae). Photo by ZHONG Hong-Tao. F, Sand-binding roots of Actinocephalus cabralensis (Ericaulaceae)(Oliveira et al., 2015). G, Dauciform roots of Tetraria species (Cyperaceae). Reprinted with permission of Elsevier B.V. (Lambers et al., 2008). H, Vellozioid roots of Barbacenia tomentosa (Velloziaceae) collected in field (Abrahão et al., 2020). EMF, ericoid mycorrhizal fungus; RE, root epidermis; RVS, root vascular system.
图3 植物养分捕获策略(NASs)随成土年龄的变化。AM, 丛枝菌根; CR, 排根; DR, 胡萝卜状根; ECM, 外生菌根; FR, 细根; N fixation, 固氮; OM, 兰花菌根; Ptase, 磷酸酶; SR, 固沙根; VR, 毛状根。背景色的深浅代表植物NASs多样性的高低。图示顶部不规则图形的宽度大小表示3种NASs随土壤年龄的变化趋势。图中圆圈大小代表不同时期NASs对植物捕获养分的相对重要性, 虚线圈代表细根策略的相对重要性还缺少足够的证据。土壤氮(N)和磷(P)库颜色的深浅代表土壤总N和P库的大小。
Fig. 3 Temporal changes in plant nutrient-acquisition strategies (NASs) with increasing soil age. AM, arbuscular mycorrhiza; CR, cluster roots; DR, dauciform roots; ECM, ectomycorrhiza; FR, fine roots; N fixation, nitrogen fixation; OM, orchid mycorrhiza; Ptase, phosphatases; SR, sand-banding roots; VR, vellozioid roots. The change in intensity of the background color represents abundance in diversity of NASs. The width of irregular polygons on the top of the figure represents the general trend of three classes of NASs with soil age. The size of cycles represents the relative importance of a particular NAS in plant acquiring nutrients in different stages of pedogenesis. The dotted circle of FR indicates that more evidence is needed to support the role of the fine root strategy in plant acquiring nutrients. The change in intensity of color in soil nitrogen (N) and phosphorus (P) pool rectangles represents variation in total N and P availability, respectively, with soil age.
图4 不同成土时期植物养分捕获策略(NASs)对植被原生演替、物种共存和多样性影响的概念图。土壤氮(N)和磷(P)库条带颜色深浅分别代表土壤N和P库的大小随成土时期的变化。成土早期第一阶段, 植物NASs主要通过促进作用影响先锋植物群落的建立。例如, 固N和释放有机酸策略提升土壤有效养分库, 促进后续物种的定居、生长和繁殖。成土早-中期, 土壤中具有较为充足的有效养分, 植物NASs主要对植物的竞争能力产生影响。具有更高效NASs的植物在竞争中具有更强的优势。因此导致物种更替, 促进演替的进行。成土后期, 土壤中的有效养分(尤其是P)几乎耗尽, 植物须发展多种互补的NASs以获取有限的养分。这些NASs间以互补关系为主。多样且互补的NASs对成土后期植物共存和多样性有重要影响, 但我们还不清楚其中的作用机理。此外, 正如Lambers等(2018)提到的, 不同NASs的植物抵御病原体的能力有差异, 也就进一步导致了这个阶段植物共存。
Fig. 4 Conceptual scheme showing effects of plant nutrient-acquisition strategies (NASs) on vegetation primary succession and coexistence and diversity of species. The change in intensity of color in soil nitrogen (N) and phosphorus (P) pool rectangle represents variations in total N and P pool with soil age, respectively. In the first stage of pedogenesis, plant NASs contribute to the establishment of pioneer community through facilitation. For example, N fixation and carboxylate-releasing strategies improve soil available nutrient pools and thus facilitate the settlement, growth and reproduction of subsequent species. In the early-intermediate stage, there are adequate soil available nutrients, NASs contribute to the competitiveness of plant species. Species with more effective NASs may have advantages in competition. And thus this will result in species turnover and promote succession. In the late stage, soil available nutrients are low or impoverished, especially for P, plant have to develop diverse NASs to acquire limited nutrients. The relationships between these NASs are mainly complementary. Diverse and complementary NASs have important impacts on the coexistence and diversity in the late stage of pedogenesis, although we still do not know the exact contribution to them. In addition, as proposed by Lambers et al. (2018), the different abilities against pathogens of plant species with different NASs also contribute the coexistence of plant species at this stage.
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