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ROGER STANIER: DIVERSITY AS THE KEY TO A NEW ERA FOR BIOLOGY JOSEPHINE ACCAPVTO-GENDRON and MORRIS GOLDNER* Introduction Roger Stanier was a profound scholar whose studies of many years concerned the basic tenets of evolution and adaptation. This Canadianborn scientist embarked on the study of microorganisms over fifty years ago and spent his working life as a microbiologist in the United States and, later, in France [I]. From his early days, taxonomy presented a constant challenge, and Stanier's interest in the subject drew him into the Bergey's Manual Trust during the years of its gestation. The Bergey 's Manual developed into the scholarly authority for determination of species. Stanier did much for its character and content, and this gave him a wide view of microbiology. His interests evolved on a broad scale from the beginning and remained so until his death some ten years ago [2]. He inspired biology students the world over through his book The Microbial World, now in its fifth edition, which he wrote with exceptional collaborators [3-7]. The depth and breadth of his studies excited research colleagues, and his ideas shaped modern biology [1, 2, 8]. The quality of his thinking was impressive; his brilliant effort to bridge the gaps in our knowledge of complex evolutionary development helped us grasp the distinctiveness of the microbial world, and his work exerts an immense and enduring influence [9]. Orientation Stanier's approach was highly influential because he perceived the study of microorganisms as a central theme of biology, one that should The authors express appreciation to Jean-Philippe Carlier, Institut of Paris for his special advice. ?University of Toronto and Institut Pasteur. (Correspondence: Department de microbiologie , Faculté de médecine, Université Laval, Quebec, Canada GlK 7P4.)© 1993 by The University of Chicago. All rights reserved. 0031-5982/93/3701-0843$01.00 48 fosephine Accaputo-Gendron and Morris Goldner ¦ Roger Stanier be in the mainstream of progress. He chose to study microorganisms in the same context as other life forms [3]. Therefore, the difficult questions that confronted biology—taxonomy, organization, inheritance, structure, function, economy, survival, evolution and adaptation— should be addressed at all levels of the natural world, including microorganisms . Stanier emphasized evolution and adaptation, explaining the control of metabolic pathways in organisms in terms of economy and adaptation for survival [10—14]. He felt the organic environment consisted of both systematic and fluctuating occurrences, thus enabling organisms to move towards successive adaptation [12]. Chatton had earlier drawn attention to the "taxonomic dichotomy" of organisms, dividing them into prokaryotes (elementary forms) and eukaryotes (complex forms) [15]. The ever perceptive Stanier worked to establish a definitive picture of prokaryotes while he assessed the major evolutionary discontinuity that separates prokaryotes and eukaryotes [3, 4, 12, 13]. However, Stanier realized that his emphasis on evolution and adaptation meant that the focus of his scientific activities should change and diversify, shifting away from taxonomy and toward biochemical facets [12]. Essentially, Stanier's work recognized the vital importance of metabolic diversities in the ever-changing environment. Before long he invoked the necessity for control systems as stable characters which would tend to be conserved with organisms [16—18]. He understood that the key to change was the availability of energy and diversity of mechanisms for its generation. Then, he was able to extend this thinking to the organisms that evolved beyond prokaryotes. He envisaged that in their evolutionary development these organisms channeled their food and energy into an ingesting activity referred to as endocytosis; he noted that endocytosis does not exist in contemporary prokaryotic forms. By depicting the course of evolution in terms of efficient endocytosis, Stanier directed attention to organellar structures and their eventual symbiotic relationships. Then, one could envision a clear understanding of these remote processes serving as a basis for the Margulis theory of the symbiotic origin of eukaryotic organelles [19—21]. Stanier explained that the variety of cytoplasmic structures bearing color pigments really reflected ancient evolutionary diversity at the prokaryotoic level [13]. Today, electron transport of the mitochondrion, an energy-generating organelle in developed eukaryotes, has an air of uniformity when compared to the diverse electron transport systems of prokaryotes for the generation of energy...


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