DEFINING COMPARATIVE ANATOMY

The title of this course might be more adequately be called "Comparative

Vertebrate Functional Anatomy".

Why emphasis on function? Integration of form and function is a fundamental

necessity in studying an organism. We need to realize that anatomy of an animal is
a certain way because an organism needs to function. The most successful
structures are those that are going to be able to persist in the population; less
successful structures will die out with their owners.

Example - Activity of wing muscles of the bird and the bat. Dissection of these
animals can give an idea of what the muscular and skeletal structures look like, but
you still need to correspond the structures to what actually goes on during flight.

What we will see this semester are variations on themes - all organisms must be
able to perform certain functions to survive. They all must perform locomotion (or
not, in the case of sessile organisms), respiration, circulation, excretion, digestion,
and reproduction. The diversity of life that we see is a product of animals finding
different ways of performing these functions.

A BRIEF HISTORY OF ANATOMY

Anatomy comes from the Greek word that means "to cut up" or "dissect"

Much of the early work in anatomy was based mostly on descriptions of organ
systems, muscle systems, and usually was conducted on domestic animals such as
livestock and chickens.

Aristotle - 4th century BC

Made detailed observations of animal anatomy. For example, he

described watching the heart of a chick develop from the liquid mass
of an egg.

Also established groupings of animals based on their structural form,

which is now known as the field of taxonomy

Galen - 2nd century AD

� Greek physician known for his comparative study of animals

� Carried out experiments to understand kidney function, movement

of blood through arteries.
 � Book on Anatomical Preparations was accepted for nearly 1400
years as the Western World�s most authoritative reference on
medical anatomy

� Downside to Galen�s work - he had little concern for pain and

suffering of his animal subjects and often dissected and examined
animals while they were still alive.

Very few advancements in anatomical study during the Middle Ages, primarily
because advancements in biological thought were also relatively infrequent during
this time.

After the Middle Ages, work moved to include the study of functional anatomy, or
the study of how structures within organisms, such as cells, tissues, organs, organ
systems, and other complex functional units, perform specific functions.

Leonardo da Vinci (15th century AD)

� His studies of anatomy, design and mechanics are well-known, as

are his sketches and work on the anatomy of flight.
William Harvey (17th century)
� Studied circulation of blood in the body, including the function of
valves in the veins from the limbs.
Giovanni Borelli (17th century)
� Applied the concept of levers (originally studied by Archimedes)
to the locomotion of animals
Carl von Linne (Carolus Linnaeus; 1707 - 1778)
� Devised the binomial system for naming plants and animal which
forms the basis of modern taxonomy

� Philosophically argued that species were unchangeable, created

originally as we find them today (based on creation as described in
Genesis)

Jean-Baptiste de Lamarck (19th century)

� Philosophie Zoologique (1809) spoke to three issues of evolution:
1) fact: species change through time - the simplest arise
through spontaneous generation from inanimate matter but
thereafter evolved onward and upward into higher forms

2) course: progressive changes in species along an ascending

scale, from the lowest/simplest to most complex/"perfect"
(humans)
 3) mechanism: need itself produces heritable evolutionary
changes - when environments or behaviors changed, an animal
developed new needs to meet the demands the environment
placed on it

* Summarized as: "Evolution by means of the inheritance of acquired

characteristics"
Georges Cuvier (19th century)
� Compared organs of various vertebrates and studied functional
relationships among the organs

� Extremely knowledgeable in the skeletal structure of animals, and

could infer the shapes of bones that would connect to neighboring
bones.

Karl Ernst von Baer, Ernst Haeckel

� Both studied comparative developmental anatomy

� von Baer noted that all early vertebrate embryos look "fishlike"
and diverge anatomically as development proceeds

� Haeckel proposed the "biogenetic law", or "ontogeny recapitulates

phylogeny", which implied that during the embryonic stages
(ontogeny) all higher animals progressively change morphologically
and resemble the evolutionary stages that preceded (phylogeny). The
theory has since been re-evaluated and given less emphasis in
comparative studies.

Charles Darwin (19th century)

� In his books On the Origin of Species (1859) and The Descent of
Man (1871) he helped to establish the evolutionary basis of our
modern synthesis of comparative, functional and adaptive
morphology and anatomy

� Proposed three conditions for and mechanisms of evolutionary

change:

1) If left unchecked, members of any species will increase

naturally in number because all species posess a high
reproductive potential.

2) Competition for resources.

 3) Survival of the few - natural selection - nature weeding out
the less fit. Superior adaptations would, on average, fare better
and survive to pass on their successful adaptations.

Alfred Wallace (19th century)

� Idependently developed the concept of "survival of the fittest"
from the observation that the human population increases faster than
food to correspond with Darwin�s "survival of the few"
Comparative anatomy as an interdisciplinary field: As with many fields,
comparative anatomy must be interdisciplinary to incorporate the full range of
factors influencing animal morphology. We now find people from all fields
involved in the study, including the following fields:
Zoology - study of animals

Physiology - study of function

Histology - study of cell and tissue structure

Genetics - study of our genetic blueprint and its effects

Ecology - study of the relationship between organisms and their

environment

Developmental Biology - study of the ontogeny of individuals from

fertilization to parturition

Evolutionary biology - study of natural selection and adaptation of

organisms to their environment

Phylogeny - comparative study of evolutionary relationships between

organisms

Research focuses on a seemingly simple question - evolution of animal body plans,

and how long ago different body plans diverged. Still, two centuries of
comparative anatomy have not yielded a consensus about the separation of
different phyla.

Although we know that a swordfish, a fly, and Marilyn Monroe all have different
body plans (representative of their phyla), we still do not know how the
characteristics unique to each arose.

Several different fields have been involved in this study to resolve the phylogenetic
tree:

Paleontologists - look at the fossil record to come up with possible

relationships based on morphology
 Paleoecologists - try to imagine how each organism may have
functioned in the environment, either as a filter feeder, predator or
grazer

Developmental biologists, molecular biologists and geneticists - have

looked at homeobox genes (which regulate the expression of other
genes and determine the features characteristic of each body segment)
to see how they are different, and may give some indication of where
each body plan diverged.

Without the cooperative efforts between these diverse fields, the exact
relationships between the different body plans might be more speculative.

This example should give you some idea of the different directions that the field of
comparative anatomy can take you.

Using comparative anatomy in the real world - medicine and beyond

At this point, you may begin thinking about how you will use this course in the
future, to get you thinking about how the concepts can be applied - most limit their
thinking of the medical applications of this class!

As a small sample of what you can potentially do:

Medicine - the techniques you learn in this class, including

memorization and integration of concepts, as well as the terminology,
will certainly help in a medical career.

Athletics and physical therapy - requires knowing how bones and

muscles interact as lever systems for maximum benefit of training
exercises as well as to minimize possibility of stress or injury to body
tissues.

Physical anthropology - understanding how stature, body shape and

limb proportions relate to environment, and apply to human origin,
distribution and ecology.

Animal behavior - how anatomical structures are used in behavior.

Examples: horns and antlers used in combat displays are related to
acquiring mating opportunities.

Evolutionary biology - anatomy helps us to understand organismal

phylogenetic relationships

Forensic anthropology - analysis of physical remains of humans to

determine identity and circumstances of death
 Biological and medical illustration

Any other areas?

MORPHOLOGICAL CONCEPTS

To analyze design, concepts of form, function, and evolution have developed

which address similarity, symmetry, and segmentation.

Similarities - corresponding parts may be considered similar to each other by:

� Homology - two or more features that share a common ancestry:

bird�s wing and mole�s arm may be traced back to common
ancestral reptile
- serial homology - special case with similarities between
successively repreated elements in the same organism:
vertebral collumn, muscle segments
� Analogy - features with a similar function: wings of bats and bees
similar in function but of different ancestral structural origin

� Homoplasy - features that simply look alike; may or may not be

homologous or analogous: turtle and dolphin flippers; insect wings
which look like leaves but cannot photosynthesize

Symmetry - how the body meets the surrounding environment:

� radial symmetry - the body is laid out equally from a central axis;
any of several planes passing through the center divids the animal
into equal halves

� bilateral symmetry - only the midsagittal section divides the body

into two equal halves

� body regions are described by basic terms of:

- anterior = head end (cranial/superior)

- posterior = tail (caudal/inferior)

- dorsal = back

- ventral = front

- the midline is medial; the sides lateral

- attached appendages have a distal (farther away) and

proximal (closer) portion
 - the pectoral region or chest supports the forelimbs

- the pelvis region refers to the hips which support the

hindlimbs

- a frontal plane divides the body into dorsal and ventral

sections, sagittal plane into left and right, and transverse plane
into anterior and posterior portions

Segmentation - a body built of repeated or duplicated segments (metameres)

separated by a series of septa.
In addition to these three, Cephalization is the pronounced tendency for the
anterior end of the body to become more and more distinctly separated and
differentiated from the rest of the body as a head.

During cephalization, the brain and sense organs become centralized at the head,
and there forms a greater elaboration of the feeding apparatus, which includes
jaws, musculature, teeth, beaks, tongues and glands.

EVOLUTIONARY MORPHOLOGY
Evolution and morphology have not always been happy companions - cooperation
between disciplines has led to concepts of design and change in design.

The concept of function covers both how a part works and how it serves adaptively
in the environment - cheek muscles of a mouse function both within an organism
(chewing) and by meeting environmental demands (resource processing), which
are defined by:

� function: the action or property of a part as it works in an organism

� biological role: how the part is used in the environment during the
course of the organism�s life history

Preadaptation: a structure or behavior posesses the necessary form and

function before the biological role arises that it eventually serves - feathers
in birds probably served as insulation to conserve body heat prior to
development of flight (thermoregulation now a secondary function)

Evolutionary change involves continuous renovations - old parts are altered

but new parts rarely added

Comparisons among characters require careful use of terminology defining

relationships - traits may be Primitive/Generalized/Derived/Specialized
An important distinction to make is among the terms. They are not necessarily
interchangeable, and should be used carefully when describing morphology.

Primitive and derived are antonyms -

� Primitive - structures that are similar to that of the ancestors or

shared by all living groups

� Derived - structures that are different from that of the ancestors

as are generalized and specialized

� Generalized - modified to perform a variety of functions

� Specialized - modified to perform restricted functions

As an example:

In mammals, the pentadactyl (five phalanges) condition is primitive, in that it is

found in all living groups. However, there is a derived condition in some
mammals, such as the bat wing, in which the first digit is elongated, or in the horse
foot, which is reduced completely to a single digit.

In contrast, our anterior phalanges (fingers) are generalized, in that they can
perform a number of different functions, from playing the piano to carving a
sculpture. However, our posterior phalanges (toes) are specialized, and can usually
only perform the function of balance and walking.

PHYLOGENY
The course of evolution (phylogeny) is often summarized in dendrograms
(schematic diagrams) that depict treelike branched connections between groups

Phylogenies serve as a graphical representation of the evolutionary relationships of

organisms. They may show:

� which organisms branched off first from a common ancestor; i.e.

the major stages of evolution of fishes (Fig. 3.6 and 3.13 in text)

� may also give information on the relative abundance of these taxa;

i.e. Fig. 1.24 in text.

Each branch in the dichotomous branching pattern signifies a point at which two
taxa diverge based on some morphological or other character trait.

All extant species usually listed in a line at the top. Extinct species� lines do not
meet up with those of extant species.
 PALEONTOLOGY
Vertebrate evolution was once referred to as the "Vertebrate Story" by
paleontologist Alfred Romer - unfolds across 590 million years with roughly
99.9% of all species which ever to have evolved now extinct

All that survives are their remnants, the fossils and scetchy vignettes they tell of
the structure and early history of vertebrates

Fossil remnants may include bones, teeth, eggs, small boney elements (embryos,
diet?), feces, DNA traces - fossil dating, restoration, and reconstruction lead to an
improved understanding of the past