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An organism is a complex chemical system that can respond to its environment, reproduce, grow, and develop in form and function, and maintain some measure of homeostasis. Nonetheless, it is a mistake to reduce the study of organisms to the level of chemistry, physics, or molecular biology. The system is highly integrated, and the properties of intact organisms are what determine their fitness in a changing environment, making the individual the primary unit of selection as well as the primary unit that interacts with the environment. Organismal biology seeks to describe and understand the responses of these complex biological systems to environmental challenges (both external and internal) as well as how they affect their environment. This approach works best when studies of model organisms are integrated into broader comparative investigations, over several levels of organization (populations, whole organisms, organs, tissues, cells, and genomes) and over time-frames from fractions of a second to millions of years. Organisms are the bridge between genomes and ecosystems, and between genetics and evolution. The impacts of environmental changes are reflected in the organism's structure, function, development, growth, evolution, distribution, and diversity, all being dependent upon its ability, or inability, to adapt and survive. Conversely, the organism shapes the environment in both subtle and profound ways. In short, as organisms go, so go their genes and their populations—and our world. The development of modern molecular biology, genetics, and genomics (along with other “omics”) has strengthened the interrelationships between field and laboratory in forming testable hypotheses for aspects of evolutionary change and responses to environmental change (Wake 2003). This two-way integration can, and does, arise at either end of a continuum extending from molecules to ecosystems in a highly hierarchical system. Integrative biology thus argues for a horizontal collaboration and cooperation in attacking important questions in biological research. This is more than a wave of the future; it is a valuable ongoing approach with incredible potential for the advancement of our base of knowledge in biology. Laying out “biology” in an overtly simplistic linear template based on structural and functional organization with molecules on one end and ecosystems on the other, we have a rather large middle ground. Organismal biology occupies this middle ground; hence any truly integrative study must pass through this region to construct large-scale concepts. When standing in this middle ground, one gets the sense that this region of organization is sometimes treated like a black box from which interesting jumps are made into molecules or populations, from laboratory or field perspectives. But returning to the organism is often neglected as the black box remains opaque. The black box nature of organismal biology is suggested when reviewers of grant proposals state as a weakness of proposed projects a lack of molecular genetic analyses of the specific research organisms (for a view of the importance of functional genomics for integrative physiology, see Dow 2007). Such views relegate established organismal models, which have contributed (significantly) to our body of knowledge but have not had their genomes sequenced, to second-class status in terms of funding importance. This can have a chilling effect on organismal biology, particularly for projects that are new and involve organisms outside the realm of standard model organisms. More importantly, this represents a shaping of research inside the black box by external factors, in effect, (potentially) creating a set of prerequisites for any new studies in organismal biology. This view negates an integrative approach that involves multiple regions of this horizontal organization, which rather than being linear is more anastomosing and ramifying in nature. The concept of integrative biology requires that each organizational region reach out to other regions to gain larger scale answers to important biological questions. A key point is that each region, including the black box that is organismal biology, should be able to originate interesting new avenues of research without external prerequisites. In other words, just as what happens within organismal biology is influenced by what happens in other areas, from molecular biology and genomics at one end and ecosystem and environmental biology at the other, new research also should flow outward from organismal biology to other areas. In our opinion, it is this opportunity to originate or initiate new avenues of integrative biology that needs to be strengthened for organismal biology. With the development of new molecular and genomic techniques, now both relatively inexpensive and fast, the need for organismal biology to concentrate on those organisms for which genomic information already exists has lessened to the extent that organismal biology can now point to the next organisms worthy of molecular and genomic analyses. That is, organismal biology can become more of an equal partner in originating and driving integrative research ventures. Of course, this is happening now, but all it takes to jeopardize this ability to originate new integrative projects is for an evaluative panel of a funding agency to buy into the argument that a research project is weak if there is no existing genomic database for the proposed research organisms. So how has organismal biology contributed to the broader approach of Integrative Biology? One extremely productive way is through the development and exploitation of modern model organisms. We use the term “modern” because, from an organismal standpoint, this approach relies at least in large part on the use of standard and/or functional genomics, in which organisms provide unique opportunities to specifically define gene functions, particularly those related to complex physiological processes (Strange 2000). As with most areas of scientific research, this approach is driven by “science through opportunity”. Modern model organisms have specific traits that make them well-suited for this form of integration, including, but not limited to, short life cycles, ease of molecular manipulation, and ability to define specific physiological variables of importance to other organisms. The best examples include species of Arabidopsis, Caenorhabditis, Dictyostelium, Drosophila, and Saccharomyces (Strange 2000). These modern model organisms, all rather strange and obscure to non-biologists, represent a hallmark of integrative biology, and are successfully boring useful horizontal tunnels through the black box of organismal biology on their journey along the continuum of biological organization. As such, this approach should be considered a flagship of organismal biology, one that is worthy of full support. But why should there be just one flagship? The horizontal tunnels of research on modern model organisms represent a relatively one-dimensional understanding of organismal biology. What is needed is a multi-dimensional method of investigation that incorporates, at the least, vertical tunnels as well. In reality, a three-dimensional approach is required, including integration outside the biological disciplines; we will leave it to others to argue this point (Schwenk et al. 2009.) This is the place of comparative biology. These vertical, anastomosing excursions are less likely to have in place many, if any, genomic data. To examine and explain the impact of a relatively new and acute environmental change on the ability of organisms to survive or thrive, we need a research strategy that is adaptive and agile, and does not have to wait on the construction of genomic databases. Rather, this new organismal research should serve as the nucleating agent for integrative research that dictates new directions from organismal biology to molecular and genomic science on the one hand, and to ecosystem and environmental science on the other. What if a change in an environmental condition, for example, allowed the creature from the Black Lagoon to expand from its limited ecological niche in the river systems of the deep jungle, where it terrorized natives, starlets, and cardboard scientists, to the beaches of the world where it could prey on humans who are just exploring their ability to contribute to the gene pool during Spring Break? (Fiction has to have that emotional and sensational draw, but organismal invasions can have equally drastic impacts on ecosystems.) Organismal biologists should not have to wait for genomic data to investigate how this large organism is capable of surviving in new environments, in particular, how it is capable of both aerial and aquatic respiration with neck gills of such small size (relative to overall body mass). Such an occurrence would represent a case of “science through opportunity,” and there is no reason why the research should not originate from within organismal biology. This research, in turn, would rely on molecular and genomic scientists to determine, among other things, the lineage of the beast, or if wide-scale mutations resulted from some unique environmental variable such as a nuclear accident (again, fiction at work). One can argue that the creature from the Black Lagoon is best studied within a comparative context, as well as in a more linear context so that integrative biology includes horizontal and vertical vectors in all directions within, and emanating from, the black box of organismal biology (the same integration should be appropriate from other levels of biological organization as well). Through this fictional analogy, we are arguing that more cross-linked research within organismal biology should be emphasized. Specifically, one strategy that should be elevated in importance, until equal to that of modern model organisms, is that of comparative biology. With new emphasis on adaptation of organisms to environmental challenges, organismal biology itself must be adaptive and agile enough to go beyond explanation of the larger scale evolutionary changes such as speciation and extinction, to focus on the more acute ecological changes including invasions and disappearances. And it should be equal to the other areas of study in its ability to originate and drive integrative science. Comparative biology has been practiced from the earliest days of recorded scientific descriptions (Sanford et al. 2002), and in many ways, it has always centered on “science through opportunity.” But this concept was pinpointed and elaborated by August Krogh and his colleagues and students. In his much cited (and quoted) paper, Krogh (1929, p. 202) issued his strong view: “For a large number of problems there will be some animal of choice or a few such animals on which it can be most conveniently studied.” But it is the end of this quotation that is of interest here: “I am afraid that most of them are unknown to the men for whom they were ‘created,’ and we must apply to the zoologists to find them and to lay our hands on them.” We suggest that this last statement be modified to be more applicable to the situation under discussion, to read (with apologies to Prof. Krogh): We are afraid that most of them are unknown to integrative biology and we must apply to the organismal and comparative biologists to find them and to lay their hands on them. We must reiterate that we are forwarding this stand without suggesting that we reduce the importance of, or the reliance on, modern model organisms. We are merely suggesting that we lift organismal biology, by elevating Comparative Biology to that same level of importance. This parallel need is summed up best by Wayne and Staves (1996, p. 369): “No single organism … exists that can provide easy access to the diversity of hidden mechanisms that underlie all interesting and important physiological and biochemical problems.” This means, as part and parcel to the argument, that we train our students to be more broadly aware of their disciplines as well as to the more integrative nature of modern scientific investigation. Students need to be able to recognize the exceptions (that the comparative approach reveals) to generalizations made from the study of modern model systems; appreciating and understanding that such exceptions lead to refinements, or even revolutions, in scientific research. One measure of a good scientist is the ability to recognize new organisms from which fundamental aspects of inquiry may be best studied, in line with the August Krogh Principle (Krebs 1975; see also Jorgensen 2001). As with all areas of scientific research these days, we must use a fluid interpretation here. For example, with modern techniques of functional genomics, organisms can be engineered (such as transgenics, knockouts, and even experimental evolution) to provide the same experimental advantages as those found in nature, so the Krogh Principle could extend fully into the arena of engineered organisms (Bennett 2003). To accept this flexibility in principle and in attitude, we must focus on what is important—how best to position our research to form a synthetic approach to complex questions in biology (Wake 2003). For organismal biology's role, at least two flagships are required. A third will be suggested by cross-disciplinary ties to other fields such as mathematics, physics, chemistry, and engineering (Schwenk et al. 2009). The light at the end of the horizontal tunnels through the black box of organismal biology, created by modern research on model organisms, is not sufficient to illuminate the full dimensions of organismal biology. Nor can it guide research at this level of organization to its full potential. This potential includes identifying and initiating integrative research on a range of biological questions extending from homeostasis in a changing environment to anatomical, physiological, and biochemical diversity in terms of the susceptibility, or robustness and resilience, of individual species and groups of species. We suggest that comparative biology be returned to the same level of importance as research on a “Modern Model Organism,” as a means of strengthening organismal biology and all of its subareas. We do not know what our next creature from the Black Lagoon will be, or how it will impact Earth's ecosystems. But with a little more illumination from the lantern of August Krogh, we will be better positioned to quickly address organismal reactions to our changing environment. Are we responsive now? Yes. We can just look to the fast response of research on hydrothermal vents (Van Dover and Lutz 2004) to see how we can react to a new creature, such as we might see with extraterrestrial life (Gaidos et al., 1999; Chyba and Phillips 2002). Can we better position ourselves for this agility, and are we at the point where organismal biology can take a leadership role in integrative biology? Absolutely. Should we do so? Absolutely again! As Ehrlich and Ehrlich (2008) detailed so well, humans are now the dominant animals on the planet, impacting every aspect of its biological support system. At the heart of the system is organismal biology, which needs to be understood from every angle as we shape the ecological and evolutionary future of our world.
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