Utilization strategies of inorganic nitrogen in submerged macrophytes
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摘要:
沉水植物在水体生态系统中发挥着关键作用，能通过直接和间接途径缓解水体中的氮负荷，促进湖泊生态系统的良性运转。沉水植物在无机氮的利用策略上与陆生植物差异显著，为了适应水环境，沉水植物地上和地下部分能够同时获取环境中的氮，并将其进行向上或向下的运输。为了减少高浓度氮的毒害，沉水植物地上和地下部分在氮利用中存在一定的权衡关系。沉水植物氮同化的主要场所是叶片，主要通过谷氨酰胺合成酶/谷氨酸合成酶（GS /GOGAT）循环和谷氨酸脱氢酶（GDH）途径进行氮的同化。目前针对沉水植物的相关研究还远不及陆生植物，仅涉及到生理响应层面，仍需深入探索沉水植物氮素利用的机理，开发合适的遗传转化体系，并利用基因编辑等技术对基因功能进行验证，对关键蛋白质的结构与功能展开深入研究。
Abstract:As important components in aquatic ecosystems, submerged macrophytes can alleviate nitrogen load and improve the healthy operation of ecosystems through direct and indirect ways. Nitrogen utilization strategies of submerged macrophytes differ significantly from those of terrestrial plants. Submerged macrophytes can uptake nitrogen not only from the overlying water, but also from pore water in sediments via above-ground and below-ground parts, respectively. To adapt to the various changes in nitrogen content in aquatic environments, submerged macrophytes exhibit two directions of nitrogen translocation, namely acropetal and basipetal translocation. To avoid the toxicity caused by high nitrogen concentrations, a trade-off exists in nitrogen utilization between the above- and below-ground parts of submerged macrophytes. The GS/GOGAT cycle and GDH pathway are the primary pathways for nitrogen assimilation in submerged macrophytes. Currently, research on submerged macrophytes lags far behind that on terrestrial plants. Further exploration of the mechanisms underlying nitrogen utilization in submerged macrophytes is still needed and a suitable genetic transformation system of submerged macrophytes is required. Molecular technologies such as gene editing can be used to identify gene function, which should promote further studies on the structure and function of key proteins.
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图 1 沉水植物在不同铵氮浓度条件下的吸收及运输过程（改自Xian等[46]）
箭头宽度代表植物地上及地下部分对外源铵态氮吸收及转运相对量的大小。GS：谷氨酰胺合成酶，GDH：谷氨酸脱氢酶，Glu：谷氨酸，Gln：谷氨酰胺。红色代谢途径为当前环境条件下植物铵同化的主要途径。
Figure 1. Ammonium uptake and transportation in submerged plants under different ammonium concentrations (modified from Xian et al.[46])
Arrow width represents relative amount of external ammonium uptake and transport by aboveground and underground parts of submerged macrophytes. GS: glutamine synthetase, GDH: glutamate dehydrogenase, Glu: glutamate, Gln: glutamine. Red metabolic pathways indicate primary route for ammonium assimilation under current environmental conditions.
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图 2 植物无机氮同化路径
GS：谷氨酰胺合成酶；GDH：谷氨酸脱氢酶；GOGAT：谷氨酸合成酶；Glu：谷氨酸；Gln：谷氨酰胺；2-OG：α-酮戊二酸。红色和黑色分别代表两条无机氮同化途径：黑色为GS/GOGAT循环途径，红色为GDH途径。
Figure 2. Inorganic nitrogen assimilation pathways in plants
GS: Glutamine synthetase; GDH: Glutamate dehydrogenase; GOGAT: Glutamate synthase; Glu: Glutamate; Gln: Glutamine; 2-OG: α-ketoglutarate. Red and black represent two pathways to inorganic nitrogen assimilation: black denotes GS/GOGAT cycle pathway, red denotes GDH pathway.
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图 3 不同适应能力及不同发育时期沉水植物的无机氮同化路径（改自Xian等[46, 68]和Ochieng等[69]）
A：高浓度铵态氮条件下，耐受型物种穗花狐尾藻铵同化以GDH途径为主导；B：敏感型物种光叶眼子菜铵同化以GS/GOGAT途径为主导； C：光叶眼子菜幼叶主要通过GDH途径进行铵的同化；D：成熟叶主要通过GS/GOGAT循环进行铵的同化。缩写同图2。
Figure 3. Inorganic nitrogen assimilation pathways in submerged macrophytes with different adaptive capacities and developmental stages (modified from Xian et al.[46, 68] and Ochieng et al.[69])
A: Ammonium assimilation in ammonium-tolerant species M. spicatum is dominated by GDH pathway under high ammonium concentration. B: Ammonium assimilation in ammonium-sensitive species P. lucens is dominated by GS/GOGAT pathway. C: In young leaves of P. lucens, ammonium assimilation is primarily mediated through the GDH pathway. D: In mature leaves of P. lucens, ammonium assimilation is mainly mediated through the GS/GOGAT cycle. Abbreviations are as in Fig. 2.
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