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Form and Function

A Photographic Exploration of Vertebrate Anatomy

Background What is the role of artistic representation in the pursuit of knowledge? Drawing of entire organisms and specific anatomical structures has always been crucial to understanding the natural world, despite a fundamental tension between art and science. Art is by nature subjective; the intentions, methods, and skill of an artist all affect how a work is perceived by the viewer, and what is taken away from a given piece varies from person to person. Science, on the other hand, strives for objectivity and clarity in writing so the hypotheses, methods, and conclusions of a given research piece are unambiguous. Museums constantly struggle with this balance. Exhibit designers want the viewer to be continually intellectually engaged while not being bored by drab aesthetics. Perhaps the most famous scientific illustrator ever was Ernst Haeckel, a German biologist, zoologist, professor and philosopher who lived from 1834 to 1919. Haeckel was an early proponent of Darwinism, believing that all natural diversity can be traced back to a single origin. While at the time he could not peer into the genome of organisms, he still found Darwins ideas to resonate in the forms (shapes) of the natural word. Haeckel famously (and controversially) postulated that ontogeny recapitulates phylogeny, claiming that the development of an organism mirrors the evolutionary history of the organism. Recapitulation theory is now thought to be half-true, as some, but not all ontogeny reflects phylogeny (Kalinka and Tomancak 2012). In 1904, Haeckel published Krustformen der Natur (Art Forms in Nature), which can be thought of as a formal argument for recapitulation theory and the idea that relationships between organisms can be unraveled by visual inspection. Krustformen der Natur is made up of plates of illustrations depicting closely related groups of organisms. Each drawing is precisely detailed and elegantly displayed; Haeckel did not explicitly need to state his support of Darwinism. Instead, his vdrawings speak for themselves, allowing the viewer to draw their own conclusions based on Haeckels visual evidence. Purpose This comparative anatomy project was inspired by Haeckels ability to combine subjective theory and objective presentation of evidence. Just as Haeckel presented external anatomy to highlight similarities within groups, this project aims to compare and contrast the internal anatomy of different groups to better understand phylogenetic relationships while straddling line line between art and science. The images in this project are meant to be beautiful while retaining an element of objectivity that allows for scientific learning to take place. Methods Between mid-November and early December, the heart, liver, and lungs (when present) were harvested from organisms dissected in the laboratory class of Vertebrate Structure and Evolution: the mudpuppy (Necturus maculosus), dogfish (Squalia acanthias), cat (Felis domestica), and pigeon (Columba livia). These four organisms each belong to separate phylogenetic groups: Amphibia, Chondrichthyes, Mammalia, and Aves. Over the course of one four-hour session, each organ was photographed with a dSLR camera separately against a white background. Ventral, dorsal, lateral, posterior, and anterior angles were all captured. Organs small enough to be spun on a single pin were photographed to get a full 360view. This included all of the hearts, as well as the synapsid livers and the pigeon lung. Finally, a 3D Model was produced using the Autodesk 123d iPhone application. The next day all of the images were imported to a computer. With Adobe Lightroom, the colors were enhanced and the pins removed from view. Using pictures of organs next to the specimens they were taken from, the organs were scaled to accurately reflect their size relative to the organism. Since the necturus lungs were dried up at the point of photographic capture, the images were omitted from the final diagrams. Columba livia

Felis domestica

Necturus maculosus

Squalia acanthias

The Liver (ventral view, scaled proportional to body length)

The liver is a vital organ in all vertebrates. While the form and function differs somewhat between vertebrates, all livers aid in the metabolism and storage of food. A diversely shaped organ across organisms, its function and shape varies widely. The squalia liver is very oily and soft, and has two posterolaterally extending lobes that take up about 30% of the squalia body mass and the bulk of space in its body cavity. Unlike osteichthyes, which have swim bladders to maintain buoyancy, sharks like squalia relies on their liver to staying afloat (Baldridge Jr 1970). By converting food into fatty acids such as squalene, sharks are able to stay buoyant in a marine environment. The necturus liver is relatively soft, and is made up of a single lobe that extends anteroposteriorly and curves dorsally. Like squalia, necturus lacks a swim bladder and uses its liver for fat storage. Since it does not live in the water column of the open ocean, however, there is less of an emphasis on buoyancy, which may explain the less oily texture compared to squalia (Fitzpatrick 1976). Thus, the necturus liver is smaller than the squalia liver in relation to body size. The felis liver is hard and rubbery with two lobes, left and right. Unlike livers of other specimens, the felis liver is dorsoventrally short. The digestive-bile-producing gallbladder is visible on the anteroventral surface. Columba has perhaps the most compressed body of all four specimens. Like felis, its liver is hard and rubbery, made up of two lobes and is dorsoventrally short. The gizzard is deep to the left lobe. To accommodate for this, the left lobe is reduced compared to the right lobe (Wischnitzer 2006). Unlike the felis heart, the columba liver is directly dorsoposterior to the heart, which is evident by the heart-shaped indentation on the ventral side.

The Heart (ventral view, scaled proportional to body length)

The heart is a muscle is that pumps blood to and from the body and respiratory organs. In the organisms examined, the heart has at least two chambers. The felis heart has four chambers: two atria, and two ventricles. Necturus has three chambers: one ventricle, and two atria (left and right). Squalia has only two chambers: the atrium and the ventricle (De Iuliis et al. 2006). Unfortunately, while looking at the hearts from the outside it is hard to determine the internal structure and blood flow. However, the atria and ventricle of necturus and squalia hearts can easily be distinguished from each other, although the division between the left and right atria in the necturus heart cannot.

The Lungs

(ventral view, scaled proportional to body length) Lungs are respiratory organs in air-breathing organisms. In felis, the lungs are made up of seven lobes: four on the right lung, and three on the left lung to make room for the heart. Like all mammalian lungs, felis lungs exhibit tidal breathing facilitated by the diaphragm that expands vascularized alveoli where gas exchange takes place. The texture of the felis lung is quite hard and rubbery. Unlike the felis lung, columbas lung is soft and porous. Air is drawn through the lungs parabronchi unidirectionally by air sacks. Thus, unlike the felis lung, columbas lung does not inflate (Wischnitzer 2006). The necturus lung is barely visible if one is not looking closely. Mudpuppies do not use their lungs much, instead relying mostly on their prominent external gills and skin for respiration. The unimportance of necturus lungs is exhibited by the poor vascularization, small size, and low number of alveoli (Wischnitzer 2006).

Lungs Heart

Heart Liver


Ventral Views

(scaled proportional to body length) For this diagram, the outlines of the organisms were scaled to each other, and then the organs were scaled to each respective organism. There are several notable differences between organisms in this diagram. Note that that relative to body size, columba has a slightly larger heart than that of felis, which are both larger than the hearts of necturus and squalia. This is because the metabolism of endothermic organisms is higher than

that of exothermic organisms (Taigen 1983; Brand et al. 1991; Herrero and Barja 1998). Squalia has the largest liver in comparison to its body size, which makes sense due to its important role in buoyancy and storage of lipids. While the liver of necturus and squalia is quite long dorsoventrally, the liver of felis and bird is dorsoventrally short and less bendable than that of squalia and necturus. The liver of felis, unlike columba, is physically separated from the heart by a diaphragm, which facilitates the expansion of the lungs in mammals (Wischnitzer 2006). Columba, in contrast,

does not need to expand its lungs, which allows for the lungs, heart, and liver to be wedged together compactly. This configuration is adaptive for aerodynamic stability by keeping the viscera closer to the center of mass, the sternum. Furthermore, maneuverability also improves as body cavities become more compact. One final notable trend to take away from these images is that the heart is always as close as possible to the primary respiratory organs. In columba and felis, the heart is directly ventral to the lungs, and in squalia and necturus it is medial to the lungs.

Lateral Views

(scaled proportional to body length) Here one can see the effects of bipedality. In columba, the organs are upright, as it walks on two legs. Felis walks on four legs, so while its organs are approximately in the same position as columba relative to the ventral surface, the angle relative to the ground is less. In necturus and squalia, it is clear that the necturus liver has more ventral curling than the squalia liver. This is unsurprising, as squalias liver is more oily and free-flowing than the more structurally stable necturus liver.

Lungs Heart Liver

Lungs Heart Liver

Liver Heart

Liver Heart

Works Cited Cat Lateral Anatomy. Diagram. Baldridge Jr H. D. 1970. Sinking factors and average densities of Florida sharks as functions of liver buoyancy. Copeia:744-754. De Iuliis G., G. DeIuliis, and D. Pulera. 2006. The dissection of vertebrates: a laboratory manual. Access Online via Elsevier. Feduccia A. 1999. The origin and evolution of birds. Yale University Press. Fitzpatrick L. C. 1976. Life history patterns of storage and utilization of lipids for energy in amphibians. Am. Zool. 16:725-732. Haeckel E., O. Breidbach, I. Eibl-Eibesfeldt, R. P. Hartmann, M. Schons, and M. Ashdown. 1998. Art Forms in Nature: The Prints of Ernst Haeckel. Prestel Munich. Herrero A., and G. Barja. 1998. H2O2 production of heart mitochondria and aging rate are slower in canaries and parakeets than in mice: sites of free radical generation and mechanisms involved. Mech. Ageing Dev. 103:133-146. Hickman Cleveland. 1992. Pigeon Anatomy. Laboratory Studies in Integrated Principles of Zoology. Diagram. Kalinka A. T., and P. Tomancak. 2012. The evolution of early animal embryos: conservation or divergence? Trends in ecology & evolution 27:385-393. Taigen T. L. 1983. Activity metabolism of anuran amphibians: implications for the origin of endothermy. Am. Nat. :94-109. Wischnitzer S. 2006. Atlas and dissection guide for comparative anatomy. Macmillan.

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