McDonald M D, Smith C P, Walsh P J
NIEHS Marine and Freshwater Biomedical Sciences Center, Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida 33149-1098, USA.
J Membr Biol. 2006;212(2):93-107. doi: 10.1007/s00232-006-0869-5. Epub 2007 Jan 30.
This review summarizes what is currently known about urea transporters in fishes in the context of their physiology and evolution within the vertebrates. The existence of urea transporters has been investigated in red blood cells and hepatocytes of fish as well as in renal and branchial cells. Little is known about urea transport in red blood cells and hepatocytes, in fact, urea transporters are not believed to be present in the erythrocytes of elasmobranchs nor in teleost fish. What little physiological evidence there is for urea transport across fish hepatocytes is not supported by molecular evidence and could be explained by other transporters. In contrast, early findings on elasmobranch renal urea transporters were the impetus for research in other organisms. Urea transport in both the elasmobranch kidney and gill functions to retain urea within the animal against a massive concentration gradient with the environment. Information on branchial and renal urea transporters in teleost fish is recent in comparison but in teleosts urea transporters appear to function for excretion and not retention as in elasmobranchs. The presence of urea transporters in fish that produce a copious amount of urea, such as elasmobranchs and ureotelic teleosts, is reasonable. However, the existence of urea transporters in ammoniotelic fish is curious and could likely be due to their ability to manufacture urea early in life as a means to avoid ammonia toxicity. It is believed that the facilitated diffusion urea transporter (UT) gene family has undergone major evolutionary changes, likely in association with the role of urea transport in the evolution of terrestriality in the vertebrates.
本综述总结了目前在鱼类尿素转运体方面的已知信息,这些信息是在其生理学以及在脊椎动物中的进化背景下得出的。人们已经在鱼类的红细胞、肝细胞以及肾细胞和鳃细胞中对尿素转运体的存在情况进行了研究。关于红细胞和肝细胞中的尿素转运情况,人们了解得很少。事实上,人们认为尿素转运体不存在于软骨鱼类的红细胞中,也不存在于硬骨鱼类的红细胞中。关于尿素跨鱼类肝细胞转运的少量生理学证据并未得到分子证据的支持,并且可能由其他转运体来解释。相比之下,关于软骨鱼类肾尿素转运体的早期发现推动了对其他生物的研究。软骨鱼类的肾脏和鳃中的尿素转运功能是使尿素在动物体内得以保留,以对抗与外界环境的巨大浓度梯度。相比之下,关于硬骨鱼类鳃和肾尿素转运体的信息是最近才有的,但在硬骨鱼类中,尿素转运体似乎起到排泄而非保留尿素的作用,这与软骨鱼类不同。在产生大量尿素的鱼类(如软骨鱼类和排尿素硬骨鱼类)中存在尿素转运体是合理的。然而,在排氨鱼类中存在尿素转运体却很奇怪,这可能是由于它们在生命早期能够制造尿素以避免氨中毒。据信,易化扩散尿素转运体(UT)基因家族经历了重大的进化变化,这可能与尿素转运在脊椎动物陆地适应性进化中的作用有关。