Beer Sven, Bjork Mats, Hellblom Frida, Axelsson Lennart
Department of Plant Sciences, Tel Aviv University, Tel Aviv 69978, Israel.Corresponding author; email:
Botany Department, Stockholm University, SE 10691 Stockholm, Sweden.
Funct Plant Biol. 2002 Apr;29(3):349-354. doi: 10.1071/PP01185.
The mechanisms by which marine angiosperms, or seagrasses, utilize external inorganic carbon (Ci) include, in addition to uptake of CO2 formed spontaneously from HCO3: (i) extracellular carbonic anhydrasemediated conversion of HCO3 to CO2 at normal seawater pH, or in acid zones created by H extrusion, and (ii) H-driven utilization (direct uptake?) of HCO3. The latter mechanism was recently indicated for Zostera marina, Halophila stipulaceaand Ruppia maritima, and manifested itself as a sensitivity of photosynthesis to buffers, as well as a relative insensitivity to acetazolamide under buffer-free conditions, especially at high pH. Seagrasses have until recently been viewed as having Ci utilization systems that are less 'efficient' than macroalgae, and this has, for example, led to the thought that future rises in atmospheric and thus dissolved CO2 would have a stronger effect on seagrasses than on macroalgae. However, most of the experiments leading to such conclusions were carried out in the laboratory on detached leaves, and buffers were used to keep HCO3/CO2 ratios stable during Ci additions. The revelation that seagrass photosynthesis is sensitive to buffers as well as to physical perturbations, has led to new experiments in which initial pH values are set by appropriate HCO3/CO3ratios, and/or O2 measurements on leaf pieces are replaced with pulse amplitude-modulated fluorometry on whole, attached leaves, often in situ. Under such conditions, the photosynthetic responses of seagrasses to Ci match those obtained for macroalgae. Thus, the paradigm of 'inefficient' Ci utilization by seagrasses as compared with macroalgae may no longer be valid. Consequently, it seems that the generally observed high productivity of seagrass beds may have its background in very efficient, H-driven, means of HCO3 utilization.
海洋被子植物,即海草,利用外部无机碳(Ci)的机制包括,除了摄取由HCO₃自发形成的CO₂外:(i)细胞外碳酸酐酶在正常海水pH值下,或在由H⁺ 外排产生的酸性区域中将HCO₃转化为CO₂,以及(ii)H⁺驱动的HCO₃利用(直接摄取?)。最近在大叶藻、盐生喜盐草和海生川蔓藻中发现了后一种机制,其表现为光合作用对缓冲剂敏感,以及在无缓冲条件下,尤其是在高pH值时对乙酰唑胺相对不敏感。直到最近,海草一直被认为其Ci利用系统比大型藻类的“效率”低,例如,这导致了一种观点,即未来大气中以及因此溶解的CO₂的增加对海草的影响将比对大型藻类的影响更大。然而,得出这些结论的大多数实验是在实验室中对离体叶片进行的,并且在添加Ci期间使用缓冲剂来保持HCO₃/CO₂比率稳定。海草光合作用对缓冲剂以及物理扰动敏感这一发现,引发了新的实验,其中初始pH值由适当的HCO₃/CO₃²⁻ 比率设定,和/或用对完整的、附着的叶片(通常是原位)进行脉冲幅度调制荧光测定法取代对叶片切片的O₂测量。在这种条件下,海草对Ci的光合响应与大型藻类的光合响应相当。因此,与大型藻类相比海草Ci利用“效率低”的范式可能不再成立。因此,似乎普遍观察到的海草床的高生产力可能源于非常高效的、H⁺驱动的HCO₃利用方式。