New and Expanded in the Sixth Edition Remarkable and innovative research continues to enrich the discipline of vertebrate biology. Much of this is added to this new edition. We now know that the regeneration of feathers is a much more complex process than previously thought, thanks to new research.
The inductive interaction between skin dermis and epidermis deep within the feather follicle establishes a zone of cell proliferation producing the feather proper, and a patterning zone where fates of newly formed cells are established in a remarkably intricate system. Feathers evolved before birds. This means that these skin specializations addressed biological roles before they addressed flight. This new description of feathers therefore opens up a new perspective on this major evolutionary event.
This is discussed in the chapter on integument chapter 6 with new supportive illustrations. Cardiac Shunt. The hearts of living amphibians and reptiles permit a right-to-left shunting of blood, thereby bypassing a trip to the lungs, but instead blood high in CO 2 heads out directly to systemic tissues.
This cardiac shunt was thought to be important during diving, where lungs quickly become depleted of oxygen and little physiological benefit attended sending blood to the lungs. This may still be true, but new and speculative research suggests another, or an additional, explanation for the shunt.
This blood bypassing the lungs may bring CO 2 to digestive organs processing a meal and thereby increase effectiveness, especially in ectothermic vertebrates. This new insight is discussed and illustrated in the circulatory system chapter chapter I have built on the genetic section on evolution and development chapter 5 introduced in earlier editions.
This has included additional illustrations and revised accompanying text. Examples throughout show how master control genes Hox genes and developmental Turning over Chordates. New developme discussed in the previous edition, informs immediate chordate ancestors flipped over, re and ventral surfaces.
That view seems to h therefore remains the surprising basis of the c plan today. Updated and Revised. Countless changes throughout this new edition have been made some small. These changes have corrected mi updated information, and often better clarifie tion.
For this I am indebted to students, re colleagues for bringing these suggestions to Serving the Student. Features of the textbo further expanded to make its presentation m inviting.
The use of color brightens these se book. Color has also been used to better compare structures between figures in these ch feasible, within color signatures, for example, more color to the illustrations. Many illustrat revised, or relabeled to improve clarity. Scientific references are available to online, if they would like to follow up or read particular subject.
The accompanying laborat guide authored with E. Zalisko is closely cro to this textbook. In addition, selective functio ries are available, online, to provide students experience of working between the anatomy tional and evolutionary significance.
Serving Instructors. This sixth edition— updated—can serve as reference and resourc the course you put together on vertebrates. I this, resources are available to you online. T laboratories may be downloaded and used as ment your course. PowerPoint images, chapt are available online along with additional McGraw-Hill that can be used to compose laboratory presentations. This laboratory manual weaves the functional and evolutionary concepts from this textbook, Vertebrates: Comparative Anatomy, Function, Evolution, into the morphological details of the laboratory exercises.
Using icons, the laboratory manual identifies cross references to this textbook, so students can quickly move from the dissection guide to this textbook to consult the expanded treatment of function and evolution. Each chapter of the dissection guide first introduces the system, makes comparisons, and demonstrates common themes in the animal systems.
Then the written text carefully guides students through dissections, which are richly illustrated. Anatomical terms are boldfaced and concepts italicized. The dissection guide is written so that instructors have the flexibility to tailor-make the laboratory to suit their needs. Here can be found the functional laboratories, helpful in a linked laboratory if available, or helpful selectively in lecture. Endof-chapter selected references, giving students a start into the literature, are located here.
Instructors can also access printable pages of illustrations that can be used as transparency masters, lecture handouts, or incorporated into PowerPoint presentations. Licensed from some of the highest-quality science video producers in the world, these brief segments range from about five seconds to just under three minutes in length and cover all areas of general biology, from cells to ecosystems.
Cours Acknowledgments I am indebted to reviewers, students, and co have generously shared with me their su improve this edition of the textbook. My hop colleagues will see, if not their point of view, influence within this edition, and accept my s for their thoughtful suggestions and criticis special help I recognize: C.
Farmer University of Utah T. Frazzetta University of Illinois Ira F. Janis Brown University Jon M. Secor University of Alabama Tamara L. Smith Westridge School Keith W. Matthias Starck, James R. Stewart, Billie J. It is again a pleasure to work with an artist as accomplished and knowledgeable as L. Laszlo Meszoly Harvard University , who contributed beautiful new figures to this edition.
I am indebted to the patient, able, and supportive people at McGraw-Hill who were so important in bringing this revised sixth edition along.
As on earlier editions, Margaret Horn was indispensible as Developm and Sue Dillon as my favorite copy editor. I the McGraw-Hill field staff who link the su of all who helped in this revision to faculty who use it. In turn, these field reps return yo of what you do and do not like, and there improvement of this textbook, making it a in progress.
Form differs because function differs. Sharks, which are a good deal denser than water, need the upward forces provided by the extended lobe of the tail to counteract a tendency to sink. Why this difference? The homocercal tail is found in teleost fishes—salmon, tuna, trout, and the like. These fishes have a swim bladder, an air-filled sac that gives their dense bodies neutral buoyancy.
They neither sink to the bottom nor bob to the surface, so they need not struggle to keep their vertical position in the water. Sharks, however, lack swim bladders, and so tend to sink. The extended lobe of their heterocercal tail provides lift during swimming to help counteract this sinking tendency.
So, the differences in structure, homocercal versus heterocercal, are related to differences in function. Why an animal is constructed in a particular way is related to the functional requirements the part serves. Form and function are coupled. Comparison of parts highlights these differences and helps us pose a question. Functional analysis helps answer our question and gives us a better understanding of animal design. Functional morphology is the discipline that relates a structure to its function.
Comparative analysis thus deploys various methods to address different biological questions. Generally, comparative analysis is used either in a historical or a nonhistorical context. When we address historical questions, we examine elucidation of an evolutionary process.
N comparisons are usually extrapolative. For testing a few vertebrate muscles, we may dem they produce a force of 15 N newtons pe timeter of muscle fiber cross section.
Rather all vertebrate muscles, a time-consuming pro ally assume that other muscles of similar cros duce a similar force other things being discovery of force production in some muscl lated to others. In medicine, the comparat drugs on rabbits or mice are extrapolated to in humans. Of course, the assumed simi which an extrapolation is based often do no analysis. Insight into the human female repro is best obtained if we compare the human cyc in higher primates because primate reprodu including the human one, differ significantly other mammals.
Extrapolation allows us to make testabl Where tests do not support an extrapolation, s served because this forces us to reflect on the behind the comparison, perhaps to reexamin analysis of structures and to return with impro ses about the animals or systems of interest itself is not just a quick and easy device.
The po size is this: Comparison is a tool of insight th analysis and helps us set up hypotheses about th mal design. Designs of Students Such philosophical niceties, however, usually students into their first course in morphology. Customarily courses prepare students headed into technica human medicine, dentistry, or veterinary med tion, morphology is important to taxonomist structure of animals to define characters. In tur acters are used as the basis for establishing between species.
Morphology is central to evolutionary b Many scientists, in fact, would like to see devoted to the combined subject, namely, morphology. The study of morphology provides its own pleasure. It raises unique questions about structure and offers a method to address these questions.
In brief, vertebrate morphology seeks to explain vertebrate design by elucidating the reasons for and processes that produce the basic structural plan of an organism. For most scientists today, evolutionary processes explain form and function. We might hear it said that the wings of birds, tails of fishes, or hair of mammals arose for the adaptive advantages each structure provided, and so they were favored by natural selection.
Certainly this is true, but it is only a partial explanation for the presence of these respective features in bird, fish, and mammal designs. The external environment in which an animal design must serve certainly brings to bear evolutionary pressures on its survival, and thus on those anatomical features of its design that convey adaptive benefits.
Internal structure itself also affects the kinds of designs that do or do not appear in animals. No terrestrial vertebrate rolls along on wheels. No aerial vertebrate flies through the air powered by a rotary propeller. Natural selection alone cannot explain the absence of wheels in vertebrates. It is quite possible to imagine that wheels, were they to appear in certain terrestrial vertebrates, would provide considerable adaptive advantages and be strongly favored by natural selection.
In part, the explanation lies in the internal limitations of the structure itself. Rotating wheels could not be nourished through blood vessels nor innervated with nerves without quickly twisting these cords into knots. Wheels and propellers fall outside the range of structural possibility in vertebrates.
Structure itself contributes to design by the possibilities it creates; evolution contributes to design by the favored structures it preserves. We must consult both structure and evolution to understand overall design. That is why we turn to the discipline of morphology. It is one of the few modern sciences that addresses the natural unity of both structure form and function and evolution adaptation and natural selection. By wrapping these together in an integrated approach, morphology contributes a holistic analysis of the larger issues before contemporary biology.
Morphology is concerned centrally with the emergent properties of organisms that make them much more than the reduced molecules of their parts. Similarly, biology has found satisfying n nations to replace what were once assumed divine causes. Modern principles of evoluti tural biology offer a fresh approach to vert and an insight into the processes responsible that design.
Like our fellow vertebrates are products of our evolutionary past and ba plan. The study of morphology, therefore, understanding of the integrated processes th To understand the processes behind our desig stand the product, namely, humans themselv we are and what we can become. But, I am getting ahead of the story. We an easy intellectual journey in reaching the c phological concepts we seem to enjoy at the m principles were not always so obvious, the always so clear.
In fact, some issues prevalent o ago remain unresolved. The significance of und ture to the evolution of design, central to mu early in the nineteenth century, is only recent amined for its potential contribution to mod ogy.
Morphology has often been internally bese contentions between those scientists centered and those centered on evolution. To some ex damental principles of both structure and ev grown from different intellectual sources and d lectual outlooks.
To understand this, we nee the historical development of morphology. Historical Predecessors—Evolu The concept of evolution is tied to the n Darwin figure 1. Although The Origin of Species was still just a few notebooks in length and several decades away from publication, Darwin had several accomplishments behind him, including his account of The Voyage of the Beagle, a collection of scientific observations. At this time, he was also engaged to his cousin Emma Wedgwood, with whom he would live a happy married life.
Nevertheless, the idea of evolution existed long before Darwin, thanks to these Greek philosophers. The first level addresses t lution and asks if organisms change through tim tion occur? The fact that evolution has occurre established by many lines of evidence, from ge the fossil record. But this does not mean that all over evolution are comfortably settled. At the might ask: What course did evolution then take anthropologists who study human evolution us the fact that humans did evolve, but they often d times violently, over the course of that evoluti can ask: What mechanism produced this evolu third level in the evolutionary debate, Darwin m contribution.
For Darwin, natural selection wa nism of evolutionary change. Verbal scuffles over the fact, course, and evolution often become prolonged and ste opponents ask questions at different levels and ing at cross-purposes. Each of these questions tled historically as well to bring us to an unders evolutionary process. Rather, Darwin proposed the conditions for and mechanism of this evolutionary change. He proposed three conditions: First, if left unchecked, members of any species increase naturally in number because all possess a high reproductive potential.
Even slow-breeding elephants, Darwin pointed out, could increase from a pair to many millions in a few hundred years. We are not up to our rooftops in elephants, however, because as numbers increase, resources are consumed at an accelerating rate and become scarce. This brings about condition two, competition for the declining resources. In turn, competition leads to condition three, survival of the few. Linnaeus devise naming plants and animals, which is still the ern taxonomy.
He founded the Museum Zoology at Harvard University. This Swedish biologist devised a system still used today for naming organisms. He also firmly abided by and promoted the view that species do not change.
Diversity of species results. To scientists of an earlier time, however, species adaptations reflected the care exercised by the Creator. Animated by this conviction, many sought to learn about the Creator by turning to the study of what He had created.
One of the earliest to do so was the Reverend John Ray 7— , who summed up his beliefs along with his natural history Deity Collected from the Appearances of Nature Agassiz — , curator of the Museum tive Zoology at Harvard University, found mu port for his successful work to build and stock a collected the remarkable creatures that wer manifestations of the divine mind figure 1.
Most of his life, Lamarck lived on poverty. Central to his philosophy vertent confusion between physiology and ev person who begins and stays with a weight-li on a regular basis can expect to see strength muscles enlarge. With added weight, use ne therefore, big muscles appear. This physiologic limited to the exercising individual because b not passed genetically to offspring.
Charles A Schwarzenegger, and other bodybuilders do n acquired muscle tissue to their children. If t seek large muscles, they too must start from their own training program. Somatic c acquired through use cannot be inherited. L ever, would have thought otherwise. Unlike such physiological responses, evo ponses involve changes in an organism that from one generation to the next. We know to characteristics are genetically based. They ar mutation, not from somatic alterations due t metabolic need.
Acquired Characteristics b a J-B. His academic position gave him a chance to promote the idea that species change. As to the mechanism of evolution, Lamarck proposed that need itself produced heritable evolutionary change.
They mistakenly view somatic parts ar immediate needs. Environmental demands do n genetic material and directly produce herita ments to address new needs or new opportuniti ing changes muscles, not DNA. Some fishes and live in deep caves not reached by daylight. T lack eyes. Even if they return to the light, eyes Evolutionarily, the eyes are lost. Its central theme holds that evolving life has a direction beginning with the lowest organisms and evolving to the highest, progressively upward toward perfection.
Evolutionists, like Lamarck, viewed life metaphorically as ascending a ladder one rung at a time, up toward the complex and the perfected. After a spontaneous origin, organisms progressed up this metaphorical ladder or scale of nature through the course of many generations. The concept of a ladder of progress was misleading because it viewed animal evolution as internally driven in a particular direction from the early, imperfect, soft-bodied forms up toward perfected humans.
As water runs naturally downhill, descent of animals was expected to run naturally to the perfected. Simple animals were not seen as adapted in their own right but rather as springboards to a better future.
The scale of nature concept encouraged scientists to view animals as progressive improvements driven by anticipation of a better tomorrow. Unfortunately, remnants of this idea still linger in modern society. Certainly humans are perfected in the sense of being designed to meet demands, but no more so than any other organism. Moles and mosquitoes, bats and birds, earthworms and anteaters all achieve an equally perfect match of parts-to-performance-to-environmental demands.
It is not the benefits of a distant future that drive evolutionary change. Instead, the immediate demands of the current environment shape animal design. The idea of perfection rooted in Western culture is perpetuated by continued technological improvements. We bring it unnoticed, like excess intellectual baggage, into biology where it clutters our interpretation of evolutionary change.
When we use the terms lower and higher, we risk perpetuating this discredited idea of perfection. Lower animals and higher animals are not poorly designed and better designed, respectively. Lower and higher refer only to order of evolutionary appearance.
Lower animals evolved first; higher animals arose after them. Thus, to avoid any suggestion of increasing perfection, many scientists prefer to replace the terms lower and higher with the terms primitive and derived to emphasize only evolutionary sequence of appearance, early and later, respectively.
For many years, textboo harsh in their treatment of Lamarck, probably his mistakes are not acquired by modern studen is also important to give him his place in the hi tionary ideas. By arguing for change in species, La blunt the sharp antievolutionary dissent of co like Linnaeus, gave respectability to the idea of e helped prepare the intellectual environment would solve the question of the origin of specie Natural Selection The mechanism of evolution by means of nat was unveiled publically by two persons in was conceived independently by both.
One Darwin; the other was Alfred Wallace. Both the respected naturalist tradition in Victorian encouraged physicians, clergymen, and person devote time to observations of plants and an countryside.
Such interests were not seen as idle time in harmless pursuits. Despite the rea was thoughtful attention to the natural world A. Wallace Alfred Russel Wallace, born in , was 14 than Darwin figure 1.
Although following naturalist, Wallace lacked the comfortable e cumstances of most gentlemen of his day; turned to a trade for a livelihood. First he surv railroads in his native England, and eventually interest in nature, he took up the collection specimens in foreign lands to sell to museums ba search for rare plants and animals in exotic land the Amazon jungles and later to the Malay A the Far East. We know from his diaries that he by the great variety and number of species to w els introduced him.
Then I at once saw, that the ever present variability of all living things would furnish the material from which, by the mere weeding out of those less adapted to the actual conditions, the fittest alone would continue the race. There suddenly flashed upon me the idea of the survival of the fittest.
The more I thought over it, the more I became convinced that I had at length found the long-soughtfor law of nature that solved the problem of the Origin of Species. Wallace, Wallace began writing that same evening and within two days had his idea sketched out in a paper. The post was slow, so the journey took four months. Darwin w geological excursions and collected biological spe graduation, he joined as de facto naturalist of the H. Beagle over the objections of his father, w to get on with a more conventional career in the He spent nearly five years on the ship and coastal lands it visited.
The experience intelle formed him. Each island own assortment of species, some found only on lar island. Local experts could tell at sight from several islands a particular tortoise came. It seemed to have suddenly dawne that not only birds, but plant and tortoise varie tinct as well. Here then was the issue.
Was each of of tortoise or bird or plant an act of spe Although distinct, each species also was clea those on the other islands and to those on the American mainland.
To account for these sp had two serious choices. The first animal or plant washed or blown to these oceanic islands would constitute the common stock from which similar but eventually distinct species evolved. Darwin sided with a natural evolution. But Darwin needed a mechanism by which such evolutionary diversification might proceed, and at first he had none to suggest.
Two years after his return, and while in the midst of writing up his results of other studies from the Beagle, Darwin read for amusement the essay on population by Malthus, the same essay Wallace would discover years later. The significance struck Darwin immediately. If animals, like humans, outstripped food resources, then competition for scarce resources would result.
Those with favorable adaptations would fare best, and new species incorporating these favored adaptations would arise. In a moment of insight, he had solved the species problem. That was , and you would think the excitement would have set him to work on papers and lecturing. Nothing of the sort happened. In fact, four years lapsed before he wrote a first draft, which consisted of 35 pages in pencil. Two years later, he expanded the draft to over pages in ink, but he shoved it quietly into a drawer with a sum of money and a sealed letter instructing his wife to have it published if he met an untimely death.
A few close friends knew what he had proposed but most did not, including his wife with whom he otherwise enjoyed a close and loving marriage. This was Victorian England. Science and religion fit hand and glove. He wanted more time to gather evidence and write the volumes he thought it would take to make a compelling case. Darwin was dumbfounded. By coincidence, Wallace had even hit upon some of the same terminology, specifically, natural selection.
Mutual friends intervened, and much to the credit of both Wallace and Darwin, a joint paper was read in the absence of both before the Linnaean Society in London the following month, July Wallace for formulating the basic concept. Da a scientific consistency and cohesiveness to t evolution, and that is why it bears the name D Science and religion, especially in Engla tightly coupled. Darwinism challenged w explanation. Controversy was immediate, and nant backwaters, it still lingers today.
Darwin h from the fray, leaving to others the task of pub the ideas of evolution. Sides quickly formed. My lord, I am on the side of the a Despite the sometimes misguided reacti cisms stuck and Darwin knew it. One was th variation, the other the question of time. As t seemed not to be enough. If the evolutionary e envisioned were to unfold, then the Earth mu to allow time for life to diversify. From his biblical studies of who begot wh historical dates available at the time, Ussher de the first day of Creation began in B October 22, at nightfall.
A contemporary, Dr vice-chancellor at Cambridge University, esti that humans were created five days later, a morning, presumably Greenwich mean time. M date as literally accurate, or at least as indi recent origin of humans, leaving no time for e apes or angels.
A more scientific effort to age made by Lord Kelvin, who used temperatures mine shafts. This fact deceptively makes it s temperature and thus in age to its molten temp formation. The true age of the Earth is actual lion years, but unfortunately for Darwin, this w until long after his death. Critics also pointed to inheritance of v weak spot in his theory of evolution.
Darwin could see this criticism coming and devoted much space in The Origin of Species to discussing sources of variation. Today we know the answers to this paradox. Mutations in genes produce new variations.
Genes carry characteristics unaltered and without dilution from generation to generation. This mechanism of inheritance was unknown and unavailable to Darwin and Wallace when they first sought answers to the origin of species. It was probably no coincidence that the intellectual breakthroughs of both were fostered by voyages of separation from the conventional scientific climate of their day.
Certainly, study of nature was encouraged, but a ready interpretation of the diversity and order they observed awaited such naturalists. Although the biblical story of creation in Genesis was conveniently at hand and taken literally by some to supply explanations for the presence of species, there were scientific obstacles as well.
Confusion between physiological and evolutionary adaptation Lamarck , the notion of a scale of nature, the idea of fixity of species Linnaeus and others , the young age of Earth Kelvin , and the mistaken views of variation and heredity blending inheritance all differed from predictions of evolutionary events or confused the picture. It is testimony to their intellectual insight that Darwin and Wallace could see through the obstacles that defeated others.
Huxley —1 Historical Predecessors—Morphology We might expect that the study of structure and the study of evolution historically shared a cozy relationship, each supporting the other. After all, the story of evolution is written in the anatomy of its products, in the plants and animals that tangibly represent the unfolding of successive changes through time. For the most part, direct evidence of past life and its history can be read in the morphology of fossils.
By degrees, living animals preserve evidence of their phylogenetic background. It might seem then that animal anatomy would have fostered early evolutionary concepts.
For some nineteenth-century anatomists, this was true. I should have thought Georges Cuvier — spanned the French Revolution, which at first won but as lawlessness and bloodshed became more of he grew increasingly dismayed by its excesses. Cert necessarily fit together. Remove whole organism fails. Consequen boasted that given one part, he c the rest. Possible combinations were thus limited to parts that meshed harmoniously and met necessary conditions for existence; therefore, the number of ways parts could be assembled into a workable organism was predictable.
Given one part of an organism, Cuvier once boasted, he could deduce the rest of the organism. Parts of organisms, like parts of a machine, serve some purpose. Consequently, for the entire organism or machine to perform properly, the parts must harmonize.
Sharp carnivore teeth would be necessarily set in jaws suited for biting, into a skull that buttressed the jaw, on a body with claws for snaring prey, with a digestive tract for digesting meat, and so forth figure 1. Alter one part, and the structurally and functionally integrated machinery of the organism would fail.
If one part is altered, function of did not occur. During his time, ancient Egyptia humans and animals were being pilfered b armies and sent to European museums. Disse that these ancient animal mummies were stru tical to modern species. Again, this was ev change, at least to Cuvier. Today, with a more sil record at our disposal and a realization th occurred over millions of years, not just withi lennia since the time of the pharoahs, we co Cuvier.
In his day, however, the mummies we sweet pieces of evidence confirming what morphology required. Parts were adapted to functions. If a part was changed, function faile mal perished.
Thus, there was no change and of species. Out of a common ancestry, evolution pas ilar structures to perform new adaptive function opposed to evolutionary ideas, was determin alternative explanation. His answer centered a types. An archetype was a kind of biologica supposed underlying plan upon which an organ All parts arose from it.
Members of each major were constructed from the same essential, b vertebrates, for instance, were thought to sh archetype, which explained why all possessed damental parts. Specific differences were fo underlying plan by particular functional need fuzzy about why he ruled out an evolutionary but he was vigorous in promoting his idea of He even carried this idea to repeated pa same individual figure 1.
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