Brain Evolution Watermark

Evolution of the Brain:
Neuroanatomy, Development, and Paleontology
g in n t r n a ri e p L e e R g r a o g f n t e o C N
Daniel D. White, Ph.D.
University at Albany, State University of New York
CONTENTS
INTRODUCTION
A look at brain size
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BASIC NEUROANATOMY 7
The basic building blocks of the central nervous system Major subdivisions of the brain 7 11 13 Major cerebral subdivisions and their functions
HUMAN BRAIN DEVELOPMENT
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LATERALIZATION, HANDEDNESS, AND LANGUAGE SEXUAL DIMORPHISM IN THE HUMAN BRAIN
COMPARING NONHUMAN PRIMATE AND HUMAN BRAINS
The indirect or comparative neuroscientific method The direct or paleoneurological method
PRIMATE PALEONEUROLOGY HOMINID PALEONEUROLOGY
Adaptive shifts in primate evolution that impact the brain
Robust and gracile australopithecine endocast morphology Early Homo Late Homo
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Homo floresiensis
SUMMARY
GLOSSARY AND KEY WORDS QUESTIONS FOR REVIEW
SUGGESTED FURTHER READING REFERENCES CITED FIGURE TITLES AND CREDITS
PREFACE
Today, the brain is considered an organ of paramount importance for those of us who study human behavior, evolution, and health. Most people would agree that it is the organ that makes us human. It is the organ that houses our personalities, memories, and thoughts. It is essentially, us. Historically, this was not always the case though. The Egyptians were famous for scooping the brains out of the skulls of deceased pharaohs and disposing of them during the mummification process. The early Greeks didnt think so highly of the brain either. Aristotle thought that it was simply the organ that cooled the heat generated by the thinkingheart. The study of the brain has come a long way since those early days and things are moving quite fast in the brain sciences. The field is moving so fast, in fact, that it is often difficult even for professionals to keep up with all of the new discoveries. I have written this module for the introductory student of human evolution and physical anthropology. In it you will be introduced to how the human brain is put together, how it functions, and how it got to be the way it is today. By studying these basic building blocks, I hope that you will be able to better appreciate and understand the other subjects covered in your course. As you read through the text, think about what parts of your brain are working to see the words, think the thoughts, and create the questions. I hope you get as much out of reading it as I did out of writing it.
I would like to thank Drs. Jurmain, Kilgore, and Ciochon for giving me the opportunity to share my knowledge of and passion for the study of human brain evolution with such a large audience. This module has been improved greatly by the careful editorial comments of Bob and Lynn. Any mistakes that may still remain are entirely my responsibility. Thanks also go to Lin Marshall at Thomson Wadsworth who patiently and cheerily guided me through the CRC process for the first time. Finally, thank you to my mentor and friend, Dr. Dean Falk, whose research continues to inspire me to learn more about how the human brain evolved.
Dan White
ACKNOWLEDGEMENTS
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INTRODUCTION
The human species is the only species that has ever asked the question, where do I come from? It is the only species that creates symphonies, poetry, and literature. It is the only species that has developed mathematical equations and explored the moon. No other species on earth is likely to ever match these achievements. The biological structure that has enabled humans to perform these feats of intelligence is, of course, the human brain. The adult human brain is a three-pound, intricately folded structure that makes up about 2% of the bodys total mass. Although relatively small in proportion to other structures in the body, the resting brain consumes as much energy as all of the resting skeletal muscles in the body put together (Aiello and Wheeler, 1995). This energetically expensive and complex organ is built on a typically primate structural plan, but also possesses some unique qualities. Studying human brain evolution through comparison with living nonhuman primates and fossilized cranial material allows paleoanthropologists to explore the questions that intrigue us most of all. When did we first become truly human? When did we first utter the sounds that everyone would consider true language? What is it about our brain that gives us the extraordinary intellectual abilities we possess today? What did the brains of our earliest ancestors look like, and what were they capable of? A look at brain size
Absolute Brain size Humans have big brains. The average adult human brain measures approximately 1350 cubic centimeters (cm3 or cc) and contains as many as 20 billion neocortical neurons (Pakkenberg and Gundersen, 1997). Human brains are by far the largest of any extant primate, weighing about three times more than the brains of our closest living primate cousin, the chimpanzee (Stephan et al., 1981). Figure 1 shows schematically how much larger the human brain is in comparison to the brains of prosimians (strepsirhines), New World monkeys, Old World monkeys, lesser apes, and great apes. This is what we call an absolute difference in brain size. For such a simple statistic, the absolute size of the brain can tell researchers a great deal about the attributes of a particular species. For example, absolute brain size (meaning the total weight or volume of all brain structures) has been shown to predict mental flexibility in nonhuman primates (Gibson et al., 2001). For our purposes, we can consider mental flexibility as a type of intelligence. Tests of mental flexibility demonstrate the cognitive power of a species by 1
challenging individuals to make decisions in a changing test environment in the laboratory. The more flexible the individual is, the better the test score, and the higher the intelligence. Just as brain size increases from prosimians to monkeys to apes, so does the cognitive flexibility of these animals. This type of experimental research is fascinating to paleoanthropologists because it takes a statistic that we can glean from fragmentary fossil evidence and allows us to make at least general predictions about the cognitive abilities of our early ancestors. We know that cranial capacity increases during human evolution from the small australopithecine brains to larger Homo habilis, Homo erectus, and eventually to the large brains of modern Homo sapiens. Experimental evidence from nonhuman primates tells us to predict greater cognitive flexibility as well.
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Figure 1. Brain volume measurements from Stephan et al. (1981) data set. (After Rilling, 2006).
Relative brain size There are certain minimum requirements for brain size in any mammal. At the lowest level, there must be enough brainpower to keep the body in homeostasis. (Homeostasis is the bodys ability to maintain internal equilibrium even as the environment changes.) There must also be enough brain and nervous tissue to sense the outside world and to respond appropriately to the stimuli from both outside and inside. These basic functions allow an animal to feed, flee danger, and reproduce. Animals that have more brain tissue than just the basic level necessary for survival may have an advantage over animals that survive with the minimum requirements for their body size. In order to determine which animals have more brainpower than is strictly necessary, we need to know the relationship between the size of the brain and the size of the body. This is called relative brain size. There are many ways to define relative brain size but the simplest is brain size divided by body size. Mammalian bodies and brains dont scale to each other in a one to one ratio. Small-bodied animals like mice have relatively large brains for their body size, while large-bodied animals like elephants or whales have relatively small brains for their body size. We see a similar phenomenon during growth. A small-bodied baby has a much larger head in relation to its body than a large-bodied adult does. This same relationship holds true for primates. As primate body size increases, the ratio of brain size to body size decreases. This is an example of allometric scaling (See Box 1). If we were to graph the relationship between body weight and brain weight in primates, it would look like a curve that flattens out as body weight increases. In order to better visualize our data, we can straighten out the allometric curve mathematically by transforming the brain and body size measurements to logarithms. Figure 2 shows the relationship between body weight and brain weight in a sample of 45 primate species drawn from the most extensively used postmortem primate data set in existence (Stephan et al., 1988; Stephan et al., 1981). The line (of regression) that runs through the data points indicates the expected size of the brain for any particular body size. For example, if we wanted to grow a 1kg (2.2 lb) galago (bush baby, see Chapter 6) to a hypothetical 60kg (130 lb) super galago, we could predict from the line its expected brain size (about 500 cm3 or 17 ounces).
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Box 1 Scaling brains and bodies: Isometry and allometry The best way to understand the scaling principles of isometry and allometry is by visualization. The diagrams below demonstrate isometry in A and allometry in B. Isometric scaling means that the shape and proportions (size ratios) of the organism in time T1 are exactly the same as the shape and proportions in times T2 and T3. The ratio of head to body size is exactly the same although both head and body size increase in absolute size. Allometric scaling means that the shape and proportions of the organism change through time. In this case, time can mean developmental time (ontogeny) or evolutionary time (phylogeny). In B, we see that the proportion of head to body size changes as the organism gets bigger. Body size increases faster than brain size and the shape of the organism in time T1 is different from the shape of the organism in times T2 and T3. This type of allometric scaling is what occurs in human development and primate brain evolution.
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The relationship between the regression line and a data point can tell us something interesting about the brainpower of that individual or group. If we look at the diamonds that represent lemurs in Figure 2, we see that, in general, these primates fall below the line whereas the human point is well above the line. This means that lemurs have smaller brains than would be predicted for a primate of equal body size, and humans have much larger brains than would be predicted for a primate of equal body size. That is, humans are more encephalized than the lemurs. In fact, humans are more encephalized than all other primates.
Figure 2. Double logarithmic scale comparing body weight in grams to brain weight in milligrams of extant primates. The line represents a linear regression that shows the expected brain size for any particular body size desired. For example, the X marks the expected location on the line of human brain size. The fact that actual human brain size is higher than the line indicates that there was selective pressure for more brain tissue during human evolution. Notice also that, in general, prosimians fall below the line and anthropoids rise above the line. Data from (Stephan et al., 1988).
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Harry Jerison, one of the worlds foremost experts in brain evolution and intelligence, first developed what is called an encephalization quotient or EQ to compare how much a species diverges from a similar line. Jerisons line included data on a large sample of living mammals. The encephalization quotient is thus the ratio of actual brain size to predicted brain size for his mammalian data set.
Encephalization quotients can give us an idea of how much more or less brainpower a species has in comparison with other mammals. As you might expect, primates have high EQ scores with humans at the top of the range. EQ scores tend to correlate highly with ecological factors like feeding behaviors and the types of food an animal eats. For example, monkeys that eat leaves have lower EQ scores than monkeys that eat fruits or apes that are omnivorous. Fruit-eating monkeys and omnivorous apes need to work harder cognitively to get their food than monkeys who eat abundant, widely available leaves. Early members of the genus Homo appear to have had a varied diet that included all sorts of food from meat to fruits and tubers. This type of diet calls for more brainpower than a simple vegetarian diet. Brain size and reorganization in the fossil record If we look at hominid phylogeny for evidence of brain size, we see that not all hominids had such large brains. Figure 3 shows that absolute brain size has been increasing since at least 1.8-2.0 million years ago when we have the first evidence of the genus Homo. Before this time, early fossil hominids like the australopithecines had brains only slightly larger than that of an average chimpanzee. Based on absolute brain size, determined by cranial capacity in fossils, it appears that members of our own genus were more mentally agile than the australopithecines and earlier members of the hominid lineage. As Kathleen Gibson and her colleagues put it, bigger is better when it comes to brain size and cognition (Gibson et al., 2001). Is an increase in absolute and relative brain size the only explanation for the presumed boost in intelligence throughout hominid brain evolution? Ralph Holloway from Columbia University would say that a rearrangement or reorganization of brain tissue and brain circuitry is also essential in the evolution of the human brain (Holloway et al., 2004). It appears that as brains get bigger theres also a change in the way they are wired up. Bigger brains require more wiring, but more wiring can become inefficient (Hofman, 2001). In order to maintain efficiency, the brain becomes reorganized. Holloway and his colleagues would say that the human brain is not just a super-sized monkey brain. To better grasp the topic of brain reorganization in hominid brain evolution, its important to first have knowledge of basic neuroanatomy and brain development. The next section provides some of these basics.
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Figure 3. Cranial capacity (a measure of brain size) of fossil hominid specimens derived from Holloway et al. (2004) and Falk et al. (2005). Notice that brain size increases more than 3 fold during hominid evolution.
BASIC NEUROANATOMY
The human nervous system can be broken down into two main components, the central nervous system and the peripheral nervous system. The central nervous system is made up of the brain and the spinal cord, which together serve as a central processing system. The peripheral nervous system is made up of nerve fibers that extend out from the brain and spinal cord to the rest of the body. These nerves interact with the external environment through sensory organs (like the skin and eyes) to let our brains know what is happening around us (exteroception). They relay information about our movement and our location in the environment (proprioception) and they communicate with organs (over which we dont have voluntary control) such as the stomach, intestines, heart, and kidneys. The basic building blocks of the central nervous system The brain and spinal cord are made up of two basic types of cells, neurons and neuroglia. Neurons are the most basic functional unit of the nervous system and are classified into three different categories: motor, sensory, and interneurons. These cells carry electrochemical impulses, called action potentials that transmit information (for example the sight and smell of your favorite food) within the brain and to other parts of the nervous system. Neurons share characteristics with other
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cells in the body but also possess specializations that allow them to do the work of thinking, moving the body, and sensing the outside world. Neuroglia (or just glia meaning, glue) are supporting cells that allow neurons to do their job properly (Barr and Kiernan, 1993). Among other functions, these cells serve as structural scaffolding for neurons and produce the insulation (myelin) that helps neurons conduct action potentials. Neurons, like other cells in the body, possess a nucleus, cytoplasm, and a selectively porous cell membrane. However, neurons also have some special qualities that allow them to carry action potentials (or more commonly nerve impulses) throughout the brain and body. These specializations are the small branch-like processes called dendrites at each end of the cell, an enlarged cell body containing a cell nucleus, and a long tail-like extension called an axon (Figure 4).
Figure 4. The basic components of a motor neuron.
In most cases, dendrites that branch off of the cell body are the receptor sites for nerve impulses coming from adjacent neurons. If a neuron adjacent to our neuron in Figure 4 is stimulated to send a nerve impulse, special signaling chemicals called neurotransmitters are released into the space that separates the two neurons. This space, called the synaptic cleft, is filled with neurotransmitters which act like keys to locked chemical gates in our neuron. Under the right conditions, these gates are opened and the process of transmitting the nerve impulse from the cell body of one neuron through the axon and on to another adjacent neuron is set in motion. This is called a chemical synapse. (There are also electrical synapses that cause action potentials but these are much less frequent than chemical synapses.) An insulation layer that surrounds the axons of most neurons called a myelin sheath allows the nerve impulses to pass rapidly through the cell. The myelin sheath acts much like the insulation around an electrical wire and just like poorly 8
insulated electrical wires, poorly myelinated neurons are inefficient. The symptoms of the chronic disease - multiple sclerosis (MS) are caused by the destruction of the myelin sheath surrounding axons throughout the nervous system (Fox, 1990). People with MS may experience blurred or double vision, poor balance, and muscle weakening to name only a few of the diffuse and numerous malfunctions caused by this disease. There are places in the central nervous system where neuron cell bodies and their accompanying dendrites are found in high density. These areas are commonly referred to as gray matter (Figure 5). Like yin and yang, where there is gray matter there must also be white matter. White matter is made up of the long myelinated axonal extensions from neurons that tend to run together in bundles or tracts. In fact, white matter is white because the myelin sheath surrounding the axons is white. One of the best examples of white matter is the corpus callosum, which serves to relay information from one side of the brain to the other.
Figure 5. The human brain in occipital view. Left side shows the cortex (gray matter) and white matter in the cerebral hemisphere. (Modified from the Comparative Mammalian Brain Collections website http://brainmuseum.org from the University of Wisconsin-Madison Brain Collection.).
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The neocortex of the cerebrum is made of six layers of cells distinguished from each other by cell type and cell density. The term neocortex (meaning new cortex) refers to this structures relatively recent phylogenetic expansion. The
neocortex is relatively large in mammals (including primates) and makes up as much as 80% of total brain volume. Evolutionary expansion of the neocortex comes in the form of increased surface area and not increased cortical thickness (Rakic and Kornack, 2001). Its the increase in surface area of the neocortex that gives the human brain its complicated pattern of gyrification. The cell composition or cytoarchitecture of the neocortex changes from one region of the brain to another and can be distinguished in histological sections. The best known and oldest map of these cytoarchitectural zones was developed by Korbinian Brodmann in his 1909 treatise (Figure 6). These zones correspond basically to functional areas of the brain such as vision, hearing, and motor control.
Figure 6. Brodmanns 1909 cytoarchitectonic map of the human brain. The numbers in the illustration designate areas that contain different combinations of cells and cell types. These areas have also been linked to various functions, such as vision (Modified from Garey, 2006).
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Major subdivisions of the brain The brain can be divided into four major regions, the brain stem (including the medulla oblongata, the pons, and the midbrain), the diencephalon, the cerebellum, and the cerebrum (Figure 7). These regions are derived from the basic hindbrain, midbrain, forebrain pattern of all vertebrate animals. The retention of this basic pattern is considered an ancestral trait in mammals. The human brain and the brains of other mammals have also evolved some derived traits that allow them to survive in their environment. Comparative neuroanatomy indicates that within the primate lineage the cerebrum (specifically the neocortex) and its components have increased in size and changed in organization more than any other structure in the brain (Rakic and Kornack, 2001). Other regions, like the olfactory bulb, have remained relatively unchanged or diminished in relative size through time (Radinsky, 1975; Stephan et al., 1988).
Figure 7. The major subdivisions of the human brain. The curved white band is the corpus callosum that links the left and right cerebral hemispheres.
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The brain stem contains structures that make connections with the spinal cord and peripheral nervous system, as well as the cerebrum and cerebellum (Figure 7). The medulla is made up of nerve fibers from the spinal cord and bundles of neurons called nuclei transmit information to higher parts of the brain
like the cerebrum. The pons (meaning bridge) carries fibers from the medulla and spinal cord to the rest of the brain, but also links sensory information processed in the cerebrum with the cerebellum. The midbrain relays some visual and auditory information from the brain to the sense organs as well as information coming from the cerebellum (Barr and Kiernan, 1993). Your midbrain is activated when you try to watch a tennis match and track the ball going back and forth over the net. The diencephalon is located at the center of the brain and contains the thalamus and hypothalamus among other important components. The thalamus is a central relay station for information passing from the peripheral nervous system, midbrain, cerebellum, and cerebrum. Its composed of densely packed nuclei that connect to areas in the cerebral cortex that process sensory (such as visual) information. The hypothalamus produces proteins called releasing factors that stimulate the pituitary gland to release hormones such as follicle-stimulating hormone, luteinizing hormone, growth hormones, and thyroid-stimulating hormones into the blood. These hormones activate target cells throughout the body that control reproduction, growth, and metabolism (Barr and Kiernan, 1993). The cerebellum is the second largest component of the brain and is best known for its involvement in keeping the body balanced and allowing it to make smooth, coordinated movements. Think about the combination of breathing, muscle-control, and balance it takes for a major league pitcher to hurl a baseball across the plate or the agility required for a gibbon to swing itself, Tarzan-like, through the forest canopy. Without a properly functioning cerebellum, this symphony of movement would be more like a hurky-jerky cacophony. Over the past several years, neuroscientists have also linked the verbal acrobatics of humans to parts of the cerebellum. It seems that searching for the right verb with the correct conjugation also requires the processing expertise of the cerebellum. The cerebrum is the largest component of the brain and contains as many as 100 distinct cortical areas that process information and initiate interaction with the environment. The next section discusses the major subdivisions of the cerebrum in more detail.
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Figure 8. Major subdivision of the cerebrum. The frontal and parietal lobes are separated by a deep consistent central sulcus. The temporal lobe is separated from the frontal and parietal lobes by the lateral or Sylvian fissure. (A fissure is a long deep sulcus.) The occipital is separated from the parietal lobe by the parieto-occipital lobe that cannot be seen in this view.
Major cerebral subdivisions and their functions There are four major subdivisions in the cerebrum and they include the occipital, temporal, parietal, and the frontal lobes (Figure 8). Each lobe specializes in a different function and contains both primary (sensory or motor) areas and multimodal association areas (Box 2). Primary areas are cortical regions that can be differentiated from adjacent areas by their unique cytoarchitecture and/or function. Association areas surround primary areas and serve to integrate information from different senses and other external or internal stimuli. Although not totally consistent, these areas can often be delineated by the sulci (sing. sulcus; valleys) and gyri (sing. gyrus; hills) of the cerebral cortex.
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Box 2 Functional mapping of the brain How do we know what parts of the brain serve which functions? The most celebrated early evidence for localizing a function to a specific area of the brain was described by Paul Broca in the early 1860s. Broca, a French surgeon and anthropologist, autopsied the brain of a man who was both paralyzed on his right side and essentially speechless. The man, who Broca called Tan, could understand when he was spoken to but could only utter one word tan. Broca observed that an injury had destroyed the left inferior part of the frontal lobe (the inferior frontal gyrus) of Tans brain. Armed with behavioral, clinical, and anatomical evidence, Broca documented the existence of the motor language area located in the inferior frontal gyrus of the left side of the brain that now bears his name (Cooper, 2006). Today, with the aid of noninvasive advanced medical imaging technology, neuroscientists and physicians are able to expand and deepen our understanding of where functions are processed in the living, thinking brain. Cognitive neuroscientists are actively designing experiments that are used to map brain activation during motor and cognitive tests to specific regions of the brain. Of greatest importance to anthropologists are those behaviors that are either adaptive or unique to humans and nonhuman primates such as advanced problem solving skills, planning behavior, complex reward seeking, and even recognizing the faces of familiar and unfamiliar species members.
Occipital lobe The occipital lobe is most closely associated with the sense of vision. Its located at the rear, or more correctly caudal (meaning toward the tail), portion of the brain. Here we find the primary visual cortex (PVC) that receives visual stimuli from the eyes. This region of the brain is also sometimes referred to as Brodmanns area 17 (Figures 6). The PVC is delineated in primates by the lunate sulcus and is surrounded by the secondary association cortex (Brodmanns areas 18 and 19) that serves to add detail and context to the raw image received by the PVC. Much of the human visual cortex is buried deep in the interhemispheric space between the two cerebral hemispheres. This makes the human visual cortex look small on the external surface of the brain compared to other primates, but it is in fact larger in absolute volume (Stephan et al., 1988). During primate evolutionary history, its possible that the visual cortex was pushed around by increasing association areas in other parts of the cerebrum. 14
Temporal lobe The temporal lobe is located below (inferior to) the Sylvian fissure and contains the primary auditory cortex, as well as auditory association cortex. The left temporal lobe contains a number of areas devoted to language such as the planum temporale and Wernickes area (Figure 9). These regions are also represented on the right hemisphere but dont perform the same function. Most of the primary auditory cortex and all of the planum temporale are buried within the Sylvian fissure and arent visible on the exterior surface of the brain. Lesions (or pathological alterations) in the association cortex around the primary auditory cortex may lead to a bizarre condition called auditory agnosia or word deafness. A patient with auditory agnosia experiences spoken words as just random sounds or noises. Damage to Wernikes area causes both word deafness and word blindness (alexia). Patients with damage to this area can speak fluently but incoherently, producing what is described as word salad. The temporal lobe is clearly essential for integrating sound information with meaning (Falk, 2000).
Figure 9. Primary areas and regions of interest in primate and hominid brain evolution. The border of the Visual area is delineated by the lunate sulcus.
Another condition thats caused by damage to either the left or right temporal lobe is called prosopagnosia. Prosopagnosia is the inability to recognize and identify faces. A patient with prosopagnosia is able to visually perceive faces but finds it impossible to use the details of a persons face to determine his or her identity (Purves et al., 1999). From an evolutionary perspective, we can see how the function of the temporal lobe is integral to the social lives of primates. A 15
highly developed ability to recognize group members and take cues from conspecifics must have provided a strong selective pressure in the highly social primate order. Parietal lobe The parietal lobe lies caudal to the central sulcus, rostral to (in front of) the lunate sulcus, and superior to (above) the Sylvian fissure (Figure 8). It contains the primary sensory area as well as a large association area. The primary sensory area receives and interprets input from the sense organs throughout the body. The human parietal association cortex has been shown to be involved in a diverse array of functions such as attention focusing, visuomotor control, and the discrimination of size and orientation of objects (Culham and Kanwisher, 2001). This multimodal association region has increased in size throughout human evolution. The parietal lobe is a good region to look for derived characteristics during primate evolution because of the diverse functions it controls. The volume of the parietal lobe may not be much larger relative to total brain size than that of living great apes but reorganization of the internal wiring could have played a major role in the behavioral and intellectual advancements seen in the genus Homo (Semendeferi, 2001; Bruner, 2004; Holloway et al., 2004; Falk, 2006) Frontal lobe The frontal lobe is the region of the brain most often associated with an individuals personality and character. This region receives input from the outside world and from primary and association areas all over the brain. The frontal lobe is our executive director and modifies sensory and association information before its acted upon. Contained in this lobe are the primary motor cortex located rostral to the central sulcus, its adjacent premotor association areas, Brocas motor speech area, and another region of special importance to hominid brain evolution, the prefrontal cortex. The functional importance of the frontal lobe is revealed when its damaged by a tumor, surgical intervention, or more temporarily with recreational drugs. We all know that we have thoughts and emotions that we control based on the social setting in which we find ourselves. Someone with a damaged frontal lobe (particularly if the damage is bilateral) is less well able to govern the expression of thoughts and emotions that rise up in daily life. For example, people with bilaterally impaired frontal lobes, who in the past were modest and reserved, may become boastful and outrageous. These people may not be able to adjust their behavior based on whether theyre surrounded by friends or in the company of their employer. They may also lose the ability to make future plans and to adapt to
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changing environments. All of these impairments take place while relative intelligence, memory, and ability to learn remain intact (Purves et al., 1999). When we look at these functions from an evolutionary perspective, we can see how important the frontal lobes have been in the development of our species. The frontal lobes are an integral part of language, the manipulation of and the adaptation to social environments, and the ability to plan ahead. All of these behaviors and skills are highly adaptive in an ever-changing environment.
HUMAN BRAIN DEVELOPMENT
Now that weve explored the basic anatomy and function of the human brain, we turn to the topic of how the brain grows. The study of growth and development from infancy to adulthood is called ontogeny. With the aid of threedimensional MRI technology, neuroscientists have been able to provide detailed evidence of the maturation rates of different cortical regions of the brain and have even linked the development of the cortex to measures of intelligence. The more we understand about the ontogeny of human brain development, the easier it is to interpret the developmental processes of extinct hominid species. At birth, the human brain has achieved only about 24-31% of its adult volume (DeSilva and Lesnik, 2006), but its blessed with an overabundance of gray matter. The neurons in this cortical gray matter come together to form an equally abundant network of synapses. After birth the brain continues to grow at a relatively fast pace but its accompanied by a period of selective pruning as well (Feldman, 2004). The overabundance of neurons, like the sucker branches of a fruit tree, are pruned away to create an efficient (and fruitful) result. In an impressive and rare longitudinal study1, Gogtay and colleagues scanned the brains of 13 healthy children every two years for 8 to 10 years. The resulting brain development maps show the trajectory of brain maturation from childhood (around 5 years old in this case) to adulthood (age 20). The researchers hypothesized that the refinement of gray matter throughout the brain would follow the functional developmental pattern seen in children. In fact, this is exactly what they found (Gogtay et al., 2004). The first areas to begin to mature are the primary sensory cortices in the parietal lobe and the frontal and occipital poles. The primary sensory cortex is considered an ancestral cortical region and is therefore shared with the earliest common ancestor of primates. Maturation progresses in a back to front direction over the primary motor cortex and eventually, at the end of adolescence, to the prefrontal cortex. In general, association cortex matures later than primary cortex
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A longitudinal study is a type of research method that enables scientists to observe and measure the same study participants over long periods of time. This type of research is valuable but also very challenging because researchers and participants both need to be committed to the project for many years.
and this has implications for behavior. For example, a five-year-old child may be very capable, physically, of throwing a baseball, but is much less able cognitively to explain the rules of the game. Similarly, the thrill-seeking behavior of the average teenager reveals the relative lack of cortical refinement in the brains executive director, the prefrontal cortex. The prefrontal cortex and its trajectory of growth have also been related to general intelligence in humans. In another long term study of 307 children and adolescents, Shaw and colleagues showed that intelligence (determined by standard Weschler intelligence scales) and the trajectory of cortical brain growth are related (Passingham, 2006; Shaw et al., 2006). These researchers used 3- dimensional MRI scans to measure the thickness of the cortex throughout the childrens development and found that, although cortical thickness doesnt correlate with intelligence, the process of cortical growth and pruning is different in children with average, high, and superior intelligence. The area that seems most profoundly influenced by this trajectory of growth is the area most closely associated with cognition and intelligence, the prefrontal cortex. It seems that children with superior intelligence show accelerated and extended cortical growth early in development with a rapid and intense period of neuronal pruning. Moreover, from an evolutionary perspective, a developmental approach has also proven valuable in studying encephalization and neocorticalization (increase in neocortex size) during primate evolution. The development of big brains and big neocortices is a function of how many neuron-producing precursor cells are created and how much time these precursor cells have to divide and proliferate (Finlay and Darlington, 1995). The more cells produced and the longer the duration of cell division, the larger the brain and the larger the brain component. Large parts of the brain, like the neocortex, have precursor cells that divide for longer periods of time than other smaller parts, like the midbrain. Similarly, organisms with large brains have extended periods of neuron precursor cell development (Kaskan and Finlay, 2001). This developmental approach reveals the elegant way that evolution works. Small changes in developmental timing may produce significant effects in the adult form and thus produce grist for the evolutionary mill.
LATERALIZATION, HANDEDNESS, AND LANGUAGE
A strange thing happens when we pick up a pen in our right hand. A message is sent from the right hand to the spinal cord and then on to the left side of the brain. The signals actually cross over to the opposite side of the brain. The same is true for the opposite side of the body. This crossover reveals some interesting things about two unique human phenomena, handedness and language.
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Out of 100 people, only about 10 are more comfortable using their left hand to write or do other basic tasks like turning a key in a lock. The remaining 90 people are much more comfortable with their right hand. This is what scientists call functional lateralization. A great deal of research has shown that other primates have hand preferences when performing certain tasks (like reaching for food or carrying an infant), but no other species is as lateralized for handedness as humans. Nonetheless, the segregation of tasks within the brain of monkeys and apes suggests that lateralization is a very old primate characteristic. I am reminded of how lateralized I am everyday when I unlock the door of my car. In a strict sense, its possible for me to use my left hand to turn the key, but the action seems effortless with my right hand. How do we explain such clear functional lateralization in handedness? From an evolutionary perspective, there doesnt appear to be any compelling selective pressure to use one hand over another. In fact, we might even argue the opposite. Think about a switch-hitting batter in baseball. A switch hitter can choose to bat lefty or righty as a consequence of the pitchers handedness. Being comfortable in all tasks with both hands would seem to be more beneficial for survival. There must be another reason why humans have become so lateralized for hand preference. Another functional lateralization that you have already read about is the location of the language centers in the brain. Brocas and Wernickes areas are located in the left hemisphere in the vast majority of people. Language is clearly an adaptive trait in humans and must have had positive selective pressure through human evolution. Interestingly, the motor language area and the primary motor area for the right hand are quite close to each other in the left hemisphere of the brain. Perhaps these areas developed synergistically and influenced the strong lateralization effects we see today (Falk, 1987). This hypothesis may work for humans but other primates dont possess the same language skills that humans do. So why are side preferences and brain asymmetries found in other primates? It turns out that as brain size increases (and remember primates in general have big brains), lateralization also increases, in part, because of the mechanical demands of brain wiring (Falk, 2006; Hofman, 2001). In order to maintain efficient communication between the parts of the brain, the whole brain becomes modularized. The left and the right sides take on different responsibilities in order to reduce the need for excess and inefficient wiring. This also has the fortunate consequence of opening up functional space for more and varied association areas. Language areas that control comprehension and the motor component of speech are on the left, while areas that control the tone of language and other more subtle functions reside in the right hemisphere (Falk, 1992). Table 1 provides a list of functional differences between the left and right sides of the human brain.
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Table 1 Functions of the Left and Right cerebral hemispheres (Falk, 1992) Functions associated with the left Functions associated with the right hemisphere hemisphere Analytical processing Global, holistic processing Language Visuospatial skills Right hand Left hand Time sequencing Recognizing faces Tone of voice Musical ability Emotions Humor/ Metaphor
These functional asymmetries are also accompanied by morphological asymmetries that can be detected in whole brains or endocranial casts (discussed below). If we look at the shape of the human brain from above, we often see that the frontal pole (or tip of the lobe) on one side extends a bit beyond the frontal pole of the other side and is a little wider laterally (Le May, 1976). The same is true for the occipital pole. These asymmetries are called petalias. The right frontal/ left occipital petalia combination is correlated with right-handedness in humans. Left-handed people tend to be more symmetrical in this aspect of brain morphology, but are more likely to show the opposite pattern to that depicted in Figure 10. Interestingly, men tend to show a greater degree of frontal/occipital petalias and are more lateralized, in general, than women. Perhaps this is why I have such a hard time opening my car door with my left hand? If an old Neandertal male had had a car, it looks like he would have had a hard time opening the door with his left hand too (Le May, 1976).
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Figure 10. Human endocast in superior view. Arrows indicate right frontal and left occipital petalias.
SEXUAL DIMORPHISM IN THE HUMAN BRAIN
Just as we cannot say that the brains of small children are simply small versions of the adult brain, we cannot say that the brains of women are just smaller versions of the brains of men. To put a twist on a favorite old cliche, it might be more accurate to say that womens brains are from Venus and mens brains are from Mars. This is perhaps an exaggeration, but it makes the point that when studying brain anatomy, function, and evolution, biological sex matters (Cahill, 2006). Humans are a sexually dimorphic species. On average, men are larger than women and have larger brains. Statistics vary from study to study, but in one large sample the brain weight of men was approximately 1392 grams and that of women was approximately 1252 grams (Ho et al., 1980). This difference has often been attributed simply to the effects of brain to body scaling. Interestingly, Dean Falk and her colleagues have shown that in a large sample of 414 human males and 390 females , the overwhelming majority of the difference between brain weight in men and women is due to sex difference rather than simply to body size difference (Falk, 2001). At the simplest level of size, the brains of men and women differ. Hundreds of neurological experiments have also shown that males tend to excel on tests of visuospatial skills, like mental rotation of objects, map reading and maze tracing, while females tend to excel in language-oriented skills like 21
reading comprehension, writing, and word puzzles. There is, however, extensive overlap between the two. So its not unusual to see a female airline pilot or a male talk show host, but the (average) differences are real (Falk, 1987; Falk, 2004a). Underlying these functional contrasts in the brains of men and women is another anatomical difference. Men have less gray matter as a total percentage of volume than women and women have less white matter as a percentage of total volume than men. In other words, women, even though they have smaller brains, have a higher percentage of that volume dedicated to the job of processing information. Males, on the other hand, have a larger portion of brain volume dedicated to passing information back and forth between the different cortical regions (Falk, 2001; Gur et al., 1999). Lets put it this way, if you and your family are lost in the middle of the woods and you need to pick a partner to go search for help, you should probably choose a male relative. When you get back from your harrowing adventure, choose a female relative to help you write the book. When did this variation in skills and brain anatomy appear in human evolution? We can only speculate. Some researchers have promoted the manufacture and use of tools as an important selective pressure for visuospatial skills in males. Other researchers prefer the idea that males were most likely to hunt and search out mates in other groups. Still others focus on the complexity of the social group and the need to band together to be successful as a selective pressure for this pattern of functional abilities. No matter what the true answer, the more we know about the differences between male and female primates, the better informed our hypotheses will be. I should also note that although the differences between male and female brains is real, these differences in no way imply superiority or inferiority of one over the other.
COMPARING NONHUMAN PRIMATE AND HUMAN BRAINS
Scientists interested in the evolution of the human and nonhuman primate brain follow two main methods of inquiry. The first method is the indirect or comparative method of study, which uses the brains of living species of primates and compares them to modern human brains. The second method is called the direct method and uses endocranial casts (endocasts) derived from fossil crania to make comparisons with endocasts of living and extinct species. The terms endocranial and endocast refer to structures on the inside of the skull. The indirect or comparative neuroscientific method The indirect or comparative method of studying brain evolution allows scientists to examine both the external, internal (hidden), and microscopic structure of the brain in a full range of primate species. The comparative method is called indirect when applied to evolutionary studies because it uses living species to 22
determine the structure and function of the common ancestor of the groups being studied. For example, to understand something about the brain of the common ancestor of chimpanzees and humans, we would study the brains of humans, bonobos, and chimps. In this approach, basic anatomical measurements can be paired with ecological, behavioral, and experimental data about these species to build a fuller picture of how the brain is influenced by the environment. For example, the strongly delineated cortical layer IV of the visual cortex in tarsiers can be linked directly to the large eyes and nocturnal behavior of this primate (Zilles, 2005) (see Chapter 6 to see a tarsier.) Examples of functional reorganization in the brain Until quite recently, neuroscientists generally agreed that the visual cortex of monkeys, apes, and humans were essentially identical (Preuss, 2001). This makes sense if we remember that stereoscopic vision and color vision are primary functions that characterize the primate pattern and are vital to the survival of these species. By studying the cortical microanatomy (meaning the pattern and types of cells) of the visual cortex, Todd Preuss and his colleagues have shown that information passing into the visual cortex from the brains main visual relay station is distributed differently in humans than in monkeys and apes. This different distribution pattern contradicts the idea of a uniform visual cortex in all primates and raises the possibility that some selective pressure unique to human evolution influenced this alteration. The researchers hypothesize that the human ability to analyze fast moving visual stimuli may aid in the process of deciphering the rapid movements of the mouth (and body) with speech (Falk, 2006; Preuss, 2001). Katerina Semendeferi and her colleagues, in a series of groundbreaking studies, utilized 3-dimensional MRI technology to compare sulcul patterns and brain volumes in humans and great apes. Table 2 shows results from a comparative analysis of whole brain and brain component measurements (Semendeferi and Damasio, 2000). As you can see from the table, human brain size is much larger than even the gorilla, whose body size is considerably larger than the average humans. The same can be said for all of the major subdivisions of the brain. However, there is a twist. As you read earlier, the frontal lobe is the region that controls higher cognitive functions, such as decision-making, planning, and the regulation of emotions. Scientists have long suspected that the frontal lobe is, if not entirely unique to humans, larger and more complex than in any other primate species. If we look solely at absolute brain volume, this is certainly the case. However, Semendeferi and Damasio also looked at what percentage of total brain size is comprised of frontal lobe and found that great apes and humans hardly differ. The 23
range for great apes is between 34.8% and 36.3%. The percentage of frontal lobe to total brain volume in humans is 36.8%. To put it a different way, the frontal lobes of humans are the volume expected for an ape of equal brain size. The question thus arises, what is it about the human brain that makes humans so much smarter than apes? Table 2 Mean whole brain and subdivision volumes in cm3 from Semendeferi and Damasio (2000) obtained by 3-dimensional MRI technology
Whole brain volume Frontal lobe volume * Temporal lobe volume* Parietal + occipital volume * 448.8 109.1 103.6 142.6 138.8 28.1 Cerebellum volume *
Human Bonobo Chimpanzee Gorilla Orangutan Gibbon
*Left and right sides combined.
The answer may be brain reorganization. The relative volume of the frontal lobe may stay constant, but a change in the way that it connects to other parts of the brain and the way its own subdivisions are laid out may have influenced the cognitive abilities humans possess today. In fact, Semendeferi and her colleagues show that Brodmanns area 10, in the prefrontal cortex of the frontal lobe, is larger relative to brain volume and potentially better wired than the same area in great apes (Semendeferi et al., 2001). A better wired prefrontal cortex is a prefrontal cortex that interfaces smoothly, rapidly, and efficiently with other areas of the brain. This suggests to these researchers that the neural substrates supporting cognitive functions associated with this part of the cortex enlarged and became specialized during hominid evolution (Semendeferi et al., 2001: 224). The direct or paleoneurological method The only direct method for studying the brain of extinct species is through the impressions of sulci and gyri, and blood vessels on the internal walls of the cranium, and the best way to do this is to create a positive cast of the endocranial surface of the cranium. Endocasts are made by pouring liquid latex or silicone into the foramen magnum at the base of the cranium. After the casting material cures, the flexible latex is drawn back carefully through the foramen magnum and popped back into shape. The crania of small-brained species and young individuals generally reproduce more details on the endocast than large crania (Connolly,
1390.3 336.3 320.9 425.1 443.2 82.3 454.8 101.5 97.2 124.8 142.3 20.3 223.2 48.2 48.2 49.8 65.7 13.5 155.1 45.8 41.2 69.3 52 10.7
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1950). Figures 11 and 12 demonstrate the morphology that is visible on a typical human endocast.
Figure 11. A rubber copy of the human brain and an endocast of the internal cranium of the same individual. Notice that there are very few distinguishable sulci and gyri in the larger human endocasts. Also note that the base of the human cast reproduces more detail than the sides or top of the cast.
Endocasts can also be formed naturally through the fossilization process as seen in the famous Taung child from South Africa. In the case of Taung, after the flesh and brain tissue decayed away, the cranium of the three year-old australopithecine tipped over on its right side and sediment filled the cavity to form a hemi-cast of the right endocranial surface. Fortunately, the fossils discoverer, Raymond Dart, was an expert in neuroanatomy and immediately recognized that the endocast and its associated cranium were the remains of a creature that had never before been described (Dart, 1925) (see Chapter 11). The most recent and most technologically advanced method for making endocasts comes from the use of 3-dimensional computed tomography (CT). Three-dimensional CT technology allows researchers to create virtual endocasts of precious fossil crania without the fear of damaging or destroying delicate internal cranial structures in the casting process (Falk, 2004b). Virtual endocasts can be manipulated with computer software in a variety of ways to obtain morphometric data (volume, linear measurement), to virtually remove tough sedimentary matrix, and to facilitate reconstruction of partial or warped casts (Conroy et al., 2000).
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Figure 12. Lateral view of a human endocast.
Paleoneurology, the study of fossil endocasts, can sometimes be controversial because not all scientists agree about what the bumps and grooves on endocasts represent (Holloway et al., 2002; White and Falk, 1999). Similarly, fragmentary fossil crania need to be reconstructed in order to evaluate endocranial morphology, and reconstructions leave room for disagreement as well (Falk et al., 2000).
PRIMATE PALEONEUROLOGY
Paleoanthropologists are interested in the evolutionary history of the Primate Order because it allows them to track the sequence and the timing of changes over primate evolutionary history. In the case of hominid brain evolution, we are particularly interested in when and why the primate brain changed from a small, rather smooth-surfaced, and tapered appearance to the large, convoluted, and rounded shape of modern ape and human brains. The history of these changes is wrapped up in the adaptive shifts that took place during primate evolution, bringing nocturnal animals into the day and arboreal animals onto the ground.
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Adaptive shifts in primate evolution that impact the brain There are three main shifts in brain morphology that took place during primate evolution: a reduction in the relative importance of olfaction, an increase in the relative importance of vision, and an increase in the relative importance of the neocortex (Radinsky, 1975) (Figure 13). What are the ecological factors that influenced these changes in brain morphology?
Figure 13. The brains of A. Bushbaby B. Macaque C. Chimpanzee drawn to approximately the same size to eliminate the effects of body size. (In reality, the bushbaby brain is 9 times smaller than the macaque brain and 40 times smaller than the chimpanzee brain.) F=frontal lobe, OB=olfactory bulb, cb=cerebellum. Notice the size of the olfactory bulb and its position in front of the frontal cortex, the size of the frontal cortex, and the position of the cerebellum under the cerebrum in the monkey and ape. (Modified from the Comparative Mammalian Brain Collections website http://brainmuseum.org from the University of Wisconsin-Madison Brain Collection.).
Modern pottos, lorises, galagos, tarsiers, and some lemurs have retained the primitive adaptation of being active at night. These prosimians are generally small and susceptible to predators like cats and large birds. Nocturnal activity paired with cryptic coloration or super-leaping ability helps these primates avoid predators. In order to communicate with each other, especially when it comes to finding mates, nocturnal prosimians use olfactory cues like urine-scenting and other odiferous secretions. This reliance on olfaction is seen in the relatively large olfactory bulbs of prosimians (Figures 13 and 14). With the exception of the South American owl monkey, all anthropoids (monkeys, apes, and humans) are diurnal. Diurnal species rely on visual cues to navigate their world. Because of their visual acuity and color vision, primates can easily detect things like the bright coloration of a ripe fruit, or the red swelling of potential mates rump; thus they use and rely upon visual input to be successful in their environment. The ability to see in three dimensions is also a highly adaptive trait because primates spent much of their evolutionary history up in trees. Falling 27
is a major selective pressure in arboreal animals. This pressure is mitigated by visual acuity and stereoscopic vision (as well as, of course, grasping hands, feet, and sometimes, tails). The relative increase in the size and complexity of the visual cortex in primates and anthropoids, in particular, is an adaptive shift of major importance. The third adaptive shift relates to the increase in both relative and absolute size of the neocortex. The neocortex, as we have already seen, is involved in the processing of information from all of the different senses. During anthropoid brain evolution, primates with more association areas in which to process information were more likely to pass on their genes to the next generation. They were cleverer, they had greater behavioral flexibility, and they were better able to adapt to changing environments.
Figure 14. Percentage of brain volume dedicated to olfactory bulbs, visual cortex, and neocortex from the Stephan et al. (1988) data set.
Eocene (56 33 m.y.a.) The first major adaptive radiation of primates began in the Eocene epoch 55 million years ago. This epoch saw the spread of small tarsier-like (omomyid) and larger lemur-like (adapid) primates around the world. The endocranial evidence for these groups of early primates is fragmentary and scarce, yet it reveals that brain size relative to body size was small, that olfaction was still quite important, and that vision was becoming better developed.
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Similar to modern prosimians, the olfactory bulb impressions of both omomyid and adapid endocasts are relatively large and protrude beyond the frontal lobe (see Chapter 7 in your text for a modern example). The surfaces of the endocasts show only a few sulci such as the Sylvian fissure and a typically lemurlike coronolateral sulcus in the adapids (Figure 15). The coronolateral sulcus courses antero-posteriorly or front to back along the cortex. This is different from the more complex mediolateral direction seen in monkeys. This simpler sulcul pattern is, in part, due to a simpler brain anatomy but is also a product of small brain size. (Remember, smaller brains tend to have fewer sulci and gyri than larger brains, even within the same phylogenetic grade.) The estimated cranial capacity of the largest Eocene specimen is less than 11 cm3 or about a third of an ounce (Falk, 2006; Gurche, 1982; Radinsky, 1975). The frontal lobes of Eocene primates arent well developed, but the parietal and occipital regions are enlarged in comparison to other early mammals. This small but detectable increase in the visual cortex of the brain foreshadows further adaptive changes in visual cortex during the evolution of monkeys and apes.
Figure 15. A. Prosimian brain showing the direction of the coronolateral sulcus; B. New World monkey brain showing the central sulcus separating the primary sensory and primary motor areas; C. Old World monkey brain. Notice the increase in the number of sulci and the shifted direction of the central sulcus in comparison to the coronolateral sulcus of the prosimian brain.
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Oligocene (33-23 m.y.a) The best paleoneurological evidence from the Oligocene is for the species Aegytopithecus zeuxis from the Fayum in Egypt. The cranial capacity of this small monkey-sized arboreal primate is only about 30 to 34 cm3 or about an ounce and it shares some primitive characteristics with early prosimians (Radinsky, 1973). Although slightly reduced in comparison with Eocene primates, the olfactory bulbs are still relatively large and extend beyond the frontal lobe in a characteristically prosimian manner. However, the endocast also reveals some morphology that foreshadows the brain development of both monkeys and apes. Miocene (23 5 m.y.a.) The Miocene has been called the golden age of hominoids because of the diversity of species, habitats, and fossil skeletons that come from this epoch. This is also the epoch where we begin to see the divergence of the ancestors of great apes and Old World monkeys from the main trunk of the hominid family tree. The fossil specimen that best exemplifies the paleoneurological evidence of the Miocene is the endocast of Proconsul . The endocast of Proconsul africanus has been reconsidered many times since its original publication in the 1950s. Compared to Aegytopithecus, Proconsul had a much larger body size and a cranial capacity that was close to 150 cm3. The endocast is warped and fragmentary, so a definitive capacity is difficult to determine. Nonetheless, the sulcul pattern of this 18 MY old African Miocene fossil indicates that the frontal lobe had begun to expand and become more complex. The central sulcus of this endocast is transverse, the visual cortex is large, and there are additional frontal sulci that reveal a more complex neocortex (Falk, 1982). The Proconsul endocast indicates that the brain of this Miocene primate was not as derived as modern ape brains, but does follow the trend of increasing size, increasing visual cortex, and increasing neocortical complexity seen in later hominoids.
HOMINID PALEONEUROLOGY
Early fossil hominid remains from all over central, eastern, and southern Africa show that hominids were bipedal long before there was any significant increase in absolute brain size. Fossil hominids (Sahelanthropus, Orrorin, Ardipithecus) from the earliest phase of hominid evolution, between 7 and 4 million years ago, so far have not provided a great deal of paleoneurological evidence. The evidence of brain evolution in the hominid phylogeny becomes more complete about a million years later with the remains of australopithecines.
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Aside from the important 3-fold increase in size between the australopithecines and modern Homo sapiens, there are several other important trends that can be elucidated by studying fossil hominid endocasts. Endocasts are judged on their level of apelike and humanlike qualities throughout hominid phylogeny. Apelike or humanlike appearance in overall shape or sulcul pattern suggests a certain level of cognitive achievement or behavioral flexibility. When we use apelike or humanlike to describe an endocast, we simply mean that the hominid being studied shares some primitive traits with the common ancestor or derived traits with later specimens. These terms dont suggest that fossil hominids were apes or more closely related to living apes. After all, living chimpanzees have been evolving for the past 6 or 7 million years too. The shape of the human endocast is rounded and globular with the cerebellum tucked up underneath the cerebral hemispheres. The endocasts of apes are more elongated and lower, with the cerebellum extending more toward the back of the cerebrum. The shape of early hominid endocasts tends to fall in between these two morphologies. We can also examine the pattern of sulci that is reproduced on the endocast surface. Although controversial, early hominids tend to show a more apelike pattern, reminiscent of the common ancestor with chimpanzees, while later fossils classified in the genus Homo tend to show more derived sulcul patterns (Falk, 1980). To illustrate these trends, we will look at a few examples. Robust and gracile australopithecine endocast morphology Chapter 11 of your textbook outlines the basic morphological and behaviorial differences between robust and gracile australopithecines. Research over the past 20 years has also revealed paleoneurogical differences between these two groups of hominids. Typical of paleoanthropological research, the similarities and differences between the gracile and robust australopithecines have generated a great deal of scientific debate. Still, its well established in paleoanthropology that the line leading to the specialized robust australopithecines eventually became extinct. On the other hand, some researchers argue that the gracile australopithecines may be the progenitor of the human lineage (Falk, 2004a). If the gracile australopithecines are related to members of the genus Homo, we might expect to find some derived characteristics that are not shared with the robust forms. However, the paleoneurological record is up for interpretation and cannot be simply read like a book as the following examples will show.
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Taung The Taung child provides an excellent example of the kind of interpretation difficulties that can arise in paleoneurology. When Raymond Dart originally described this fossil, he mistook the impression of the lambdoid suture (the location where the parietal and occipital bones knit together) for the lunate sulcus (Falk, 2004a). The lunate sulcus, you will remember, is the depression that delineates the primary visual cortex from secondary association areas in the occipital lobe. By misinterpreting the location of this sulcus, the endocast of Taung looked to have a humanlike occipital lobe. This was, of course, tremendously important from a psychological point of view because it gave the small man-ape from South Africa an air of legitimacy in the human family tree. In a reanalysis of South African fossil hominid natural endocasts, Falk (1980) demonstrated that a small dimple in an apelike position was more likely the position of the lunate sulcus. Holloway disputed this location and preferred a rearward position that would make the gracile australopithecine occiput look more humanlike. No one will ever know the true location of the lunate sulcus in the Taung specimen, but further evidence from the front part of the endocast suggests that if Taung had a humanlike occiput, it had an apelike frontal lobe. Thus, the Taung lunate sulcus is only part of the story. The relatively complete frontal lobe of this specimen reveals both a sulcul pattern and sulcul density that is reminiscent of ape morphology (Falk, 1980). One example of this apelike morphology is the presence of a fronto-orbital sulcus that is nonexistent in human brains and shared with great ape brains (Figure 16). The fronto-orbital sulcus of a chimpanzee brain is depicted in Figure 16B. This suggests that the sulcul pattern of australopithecines had a primitive characteristic shared with the common ancestor of hominids and chimpanzees. Humans have a derived pattern of gyri and sulci caused by reorganization frontal cortices (Figure 16A).
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Figure 16. A. Human brain B. Chimpanzee brain. fi = inferior frontal sulcus, R= horizontal branch of the Sylvian sulcus, R= ascending branch of the Sylvian sulcus, S= Sylvian fissure, ls = lunate sulcus, fo = fronto-orbibital sulcus. Notice the relatively forward position of the lunate in the chimpanzee brain.
Reconstructing robust endocasts Reconstructing fragmentary fossil hominid endocasts is a tricky business. The process takes both artistic talent and an intimate knowledge of human, ape, and fossil hominid endocranial anatomy. It is therefore crucial that early reconstruction attempts are analyzed critically when new fossil hominid material is discovered. In 2000, a group of researchers did just this with robust australopithecines. Early reconstructions of robust australopithecine endocasts were apparently based largely on more complete endocranial remains from gracile australopithecines like the famous Mrs. Ples fossil, Sts 5 from Sterkfontein, and thus resembled gracile australopithecines in some ways (Falk et al., 2000). With the discovery of several new robust australopithecine remains, Falk and her colleagues showed that the frontal lobes of robust australopithecines were much different in shape than gracile australopithecines. It appears that the robust
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australopithecines have more beak-shaped (pointed) frontal lobes than the more rounded lobes of the gracile forms. As we have noted above, the frontal lobes are critical regions for advanced thinking, language, and higher cognitive abilities in humans. Rounded and expanded frontal lobes, in addition to derived characteristics in the temporal lobe in the gracile australopithecines, indicates that there was reorganization taking place in critical areas of the brain of early fossil hominids. While apelike in sulcul pattern, frontal lobe expansion in gracile australopithecines foreshadows the morphology of early Homo. Another interesting and controversial difference between the endocasts of robust and gracile australopithecines is the pattern of cranial venous sinuses in the back of the cranium. Venous sinuses are the pathways by which blood drains from the brain. Robust australopithecines have an enlarged occipital/marginal (O/M) venous drainage system that is rare in humans and detectable in only one gracile australopithecine. Instead, humans have large transverse and sigmoid sinuses (Figure 12). This divergent pattern of blood drainage has been used, in part, to help explain the increased potential for bigger brains in later hominids. The radiator hypothesis, as it is called, relates the hominid pattern of obligate bipedalism and brain size increase to the challenges of living in a hot equatorial environment. Some researchers have hypothesized that an efficient brain cooling mechanism (via venous sinuses and emissary vessels) allowed the genus Homo to benefit from continued brain size expansion and reorganization. (See Falk, 2004a for a full explanation.) Early Homo When did we first become truly human? When did we first utter the sounds that everyone would consider true language? These are fascinating questions, the answers to which will only be obtained if we invent a time machine. We can, however, look to the archeological and paleoneurological record for some interesting evidence. The fact that scientists have chosen to categorize some extinct hominid species in the genus Homo says something about the relative human-ness of these groups. The archeological evidence of more complex tools implies increased cognitive complexity in Early Homo. The paleoneurological record of Early Homo also shows a step in the direction of more humanlike morphology and perhaps, more humanlike behavior. One of the most famous fossil hominids KNM-ER 1470 (the best preserved Homo habilis skull), is represented by a reconstructed, mostly complete, cranium. The endocranial volume of ER 1470 measures 752 cm3 (which is slightly more than half the size of modern human cranial capacity), and the endocast reproduces the earliest humanlike sulcul pattern described to date, including what has been 34
described as Brocas motor speech area. It would therefore be very easy to jump to the conclusion that ER 1470 was capable of humanlike speech but this conclusion would be hasty. We dont know if Brodmanns areas 44 and 45 in humans are directly comparable to the same areas in ER 1470, so we can only cautiously state that the sulcul pattern of ER 1470 more closely resembles the pattern of modern humans than modern apes and australopithecines. Late Homo The endocasts of later hominids like Homo erectus and Homo heidelbergensis (as well as Neandertals) are large and reproduce humanlike asymmetries (petalias and Brocas caps), but tend to differ in sulcul details seen in earlier smaller-brained hominids. The reason for this variability is unknown, but it may have something to do with the thickness and durability of the brains protective coverings called the meninges. The shape of Homo erectus endocasts is reminiscent of the craniums wide base and long, low profile. The leap toward human morphology is evident in H. erectus, but the frontal lobes dont appear as round and prominent as in the modern endocast. Neandertal endocasts are large and clearly encephalized but possess uniquely protuberant occipital lobes that may reveal an expansion of visual cortex in these extinct hominids (Holloway et al., 2004). The archeological record of these later Homo species shows that these groups had migrated around the Old World, used complexly designed tools, and lived in larger, more complex groups. Large, complex neocortices with greater association capacity were clearly positive adaptations for these hominids. In fact, the size of the neocortex is highly correlated with social group size and complexity (Dunbar, 1998). Cooperation and social interaction puts a premium on efficient symbolic communication, which in turn, contributes to higher reproductive success in even the harshest environments. Homo floresiensis In October 2004 Peter Brown and his colleagues introduced to the world a new species of hominid called Homo floresiensis that survived until around 13,000 years ago, making it the latest surviving species in the genus Homo other than our own. The bones of H. floresiensis indicate an extremely small stature for a member of the genus Homo (about 3 feet tall) and an equally small cranial capacity (417 cm3) (Falk et al., 2005). The skeletal remains were also associated with a diverse group of well-made stone tools (see Chapter 14). Fortunately, among the skeletal remains was a very well-preserved cranium that enabled paleoanthropologists to create a virtual endocast using 3-dimensional CT. The endocast of the specimen called Liang Bua 1 (LB1) was compared to a 35
series of ape, human, and fossil hominid virtual endocasts. The results of this study showed that although the endocranial volume of LB1 is similar to the much more ancient australopithecines, the shape is most similar to that of Homo erectus from Asia. Like H. erectus, the endocast is wider at the back - and lower portions are long and low in profile (Falk et al., 2005). Furthermore, the pattern of sulci on the frontal, parietal, and occipital lobes appear to be humanlike in configuration. The position of the lunate sulcus is similar to that in humans, and the distinctive apelike fronto-orbital sulcus is absent. Other than its extremely small size, the endocast of LB1 is clearly quite derived. There has also been some suggestion that this individual suffered from a genetic disorder called microcephaly that leads to abnormally small brain size. Based on comparative evidence, this suggestion doesnt seem to be the best explanation for the morphology that is reproduced on the LB1 endocast. It appears normal in shape, just small in size, although debate continues (Falk et al., 2006; Falk et al., 2007; Jacob et al., 2006; Martin et al., 2006; Weber et al., 2005). This fascinating find extends the range of Homo cranial capacity from the 1600 cm3 high of Neandertals to the low 417 cm3 of H. floresiensis (Figure 3). What does this range say about the intellectual capacity of hominids from one end of the spectrum to the other? Did the hobbits from Flores use symbolic communication like modern humans? Did they use their brains to manipulate the social environment to get ahead? How much brain tissue does a species need to be considered human? The questions are easier to pose than the answers are to find or harder yet to get scientists to agree about.
SUMMARY
Evolution has acted on the brains of primates to mold a highly flexible, intelligent, and visually-oriented organism. Anthropoids (monkeys, apes, and humans) tend to diverge more than prosimians from the less-derived brain morphology of the common ancestor of primates. The brains of anthropoids reflect a clear reduction in the neural tissue allocated to olfaction, an increase in visual processing cortex, and a trend toward more complicated association areas in the neocortex. These trends are maximized in the hominid lineage. Small advances in neocortical association areas and brain wiring enabled hominids to plan ahead, work in groups, and communicate more effectively. Brainpower began to replace brute strength, and the energy demands of successful adaptive responses shifted. The uncertain and dangerous environment of early hominids created an important selective pressure on the brains of these terrestrial bipeds. Those individuals that were able to eke out a living by using tools, planning migration routes, remembering the location and season of fruiting trees, and hunting wild game were
36
more likely to pass on their genes to the next generation. Their most powerful tool was their brain. The brain is a powerful tool not only for competing with other species in an ecological niche but also for competing with members of ones own group. Social intelligence or the ability to navigate a rich and complex network of family, friends, and foes developed along with brain size during hominid evolution. When it comes to keeping track of social relationships and large brains, Homo sapiens is the species par excellence. Today, the neurocentric age is in full swing (Zimmer, 2004). We learn in leaps and bounds. Details of how we store memories, how we process thoughts, how we form sentences, and how we solve problems will all become clearer in time. New technologies will allow us to delve more deeply into the thinking brain and to look more closely at intraspecific diversity (Holloway et al., 2004). It is a very exciting time, indeed.
GLOSSARY AND KEY WORDS

Absolute brain size Action potentials Alexia
Ancestral
Association areas Auditory agnosia Axon
Brain stem Caudal
Central nervous system Chemical synapse Coronolateral sulcus
Total weight or volume of all brain structures Word blindness Retaining the same traits as the common ancestor Zones of integration of several modes of external information Word deafness Long tail-like extension from the body of the neuron The midbrain, the pons, and the medulla oblongata Toward the tail or rear The brain and the spinal cord Sulcus seen in prosimians and early primates but not 37
Electrochemical impulses more commonly known as nerve impulses
Release of neurotransmitters across the synaptic cleft that initiates an action potential
humans Cortical microanatomy Cytoarchitecture Derived The pattern and types of cells in the cortex The cell composition of the neocortex
Diencephalon
Direct method Encephalized Endocasts
Exteroception Frontal pole
Gyri (sing. Gyrus) Gyrification
Homeostasis
Indirect method Lateralization
Possessing different traits than the common ancestor Brain region that contains the thalamus and hypothalamus Studying the inside of the cranium of fossils through endocasts Having a larger brain than predicted by body size A cast (plaster, latex, silicon, or virtual) of the inside of the cranium Perceiving the external environment The most anterior (front) part of the cerebral hemispheres The hills in the complicated neocortical pattern The pattern of hills and valleys that make up the neocortex The bodys ability to maintain internal equilibrium even as the environment changes One side functions differently or has greater facility with tasks than the other side of the body. Example: handedness. Furrow or valley that delineates the primary visual 38
Comparative method of studying brains where humans are compared to other primates
Lesions Lunate sulcus
Pathological alterations in the brain caused by loss of blood flow or physical damage
cortex Myelin Neocortex Insulation around nerve cell axons Meaning new cortex, is the structure that contains six layers of cells and controls the bodys higher functions like thinking Support cells in the nervous system that provide structure and insulation to neurons
Neuroglia (glia) Neurons
Neurotransmitters Nuclei
Occipital lobe Occipital pole Ontogeny
Parietal lobe
Peripheral nervous system Petalia
Planum temporale Primary areas Proprioception
Cell that is the basic functional unit of the nervous system Special signaling chemicals released by neurons Bundles of neurons The most posterior (back) part of the cerebral hemispheres The study of growth and development Top rear part of the brain that contains essential association areas Area in the temporal lobe that is associated with language in humans Early developing areas that have control over one major modality (vision, hearing, motion, sensation) Perceiving your own body in its environment 39
Rear part of the brain that contains the primary visual cortex
Nerve fibers that extend out from the brain and spinal cord to the rest of the body The asymmetrical shape pattern of human brains. One side usually projects forward more than the other and is wider.
Prosopagnosia Relative brain size Rostral
The inability to recognize faces Brain size divided by body size Toward the front
Sulci (sing. Sulcus) Superior
Synaptic cleft
Temporal lobe
Venous sinuses
Wernickes area
QUESTIONS FOR REVIEW
The valleys in the complicated neocortical pattern Above in anatomical terms The space between the terminal ends of one neuron and another Lower lateral portion of the brain that contains the primary auditory cortex System of veins that drains blood from the brain and head 40
Area in the temporal lobe (mainly) that is involved in the comprehension of language in humans
1. What is the difference between relative and absolute brain size and what can these measurements tell us about the life and behavior of an animal? 2. What are the major components of the human brain and what basic functions are they responsible for? 3. How can studying brain development in human children help us better understand the evolution of fossil hominid brains? 4. How is language linked to brain lateralization? 5. What are some of the basic differences between male and female brains? 6. What methods do scientists use to study brain evolution? 7. In what areas do human and prosimian brains differ most? 8. What are the major adaptive shifts in primate evolution that impact the brain? 9. Why is the Taung child significant to scientists who study the brain? 10. Why is the discovery of Homo floresiensis significant for anthropologists who study the brain?
SUGGESTED FURTHER READING

Braindance: Revised and Expanded Edition by Dean Falk Soul Made Flesh by Carl Zimmer The Human Fossil Record, Volume 3, Brain Endocasts - The Paleoneurological Evidence by Ralph Holloway and colleagues.
REFERENCES CITED
Aiello L, and Wheeler P (1995) The expensive-tissue hypothesis: The brain and the digestive system in human and primate evoluton. Current Anthropology 36:199-221. Barr M, and Kiernan J (1993) The human nervous system. An anatomical viewpoint. New York: JB Lippencott.
Bruner E (2004) Geometric morphometrics and paleoneurology: brain shape evolution in the genus Homo. J Hum Evol 47:279-303.
Cahill L (2006) Why sex matters for neuroscience. Nat Rev Neurosci 7:477-84. Connolly C (1950) External anatomy of the primate brain. Springfield, Illinois: Thomas. Conroy GC, Falk D, Guyer J, Weber GW, Seidler H, and Recheis W (2000) Endocranial capacity in Sts 71 (Australopithecus africanus) by threedimensional computed tomography. Anat Rec 258:391-6.
Cooper DL (2006) Broca's arrow: evolution, prediction, and language in the brain. Anat Rec B New Anat 289:9-24. Culham JC, and Kanwisher NG (2001) Neuroimaging of cognitive functions in human parietal cortex. Curr Opin Neurobiol 11:157-63.
Dart R (1925) Australopithecus africanus: The Man-Ape of South Africa. Nature 115:195-199. DeSilva J, and Lesnik J (2006) Chimpanzee neonatal brain size: Implications for brain growth in Homo erectus. Journal of Human Evolution 51:207-212.
41
Dunbar R (1998) The social brain hypothesis. Evolutionary Anthropology 6:178109.
Falk D (1980) A reanalysis of the South African australopithecine natural endocasts. Am J Phys Anthropol 53:525-39. Falk D (1982) A reconsideration of the endocast of Proconsul africanus. In R Ciochan and R Corruccini (eds.): New interpretations of ape and human ancestry. New York: Plenum Publishing Company, pp. 239-248. Falk D (1987) Brain lateralization in primates and its evolution in hominids. Yearbook of Physical Anthropology 30:107-125.
Falk D (1992) Evolution of the brain and cognition in hominids: Sixty-Second James Arthur Lecture on th evolution of the human brain. American Museum of Natural History, New York.
Falk D (2000) Hominid brain evolution and the origins of music. In N Wallin, B Merker and S Brown (eds.): The origins of Music: MIT, pp. 197-216. Falk D (2001) The evolution of sex differences in primate brains. In KR Gibson and D Falk (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press, pp. 98 - 112. Falk D (2004a) Braindance. Gainesville: University Press of Florida.
Falk D (2004b) Hominin brain evolution--new century, new directions. Coll Antropol 28 Suppl 2:59-64.
Falk D (2006) Evolution of the Primate Brain. In W Henke, H Rothe and I Tattersall (eds.): Handbook of Paleoanthropology. Amsterdam: SpringVerlag.
Falk D, Hildebolt C, Smith K, Morwood MJ, Sutikna T, Brown P, Jatmiko, Saptomo EW, Brunsden B, and Prior F (2005) The brain of LB1, Homo floresiensis. Science 308:242-5.
Falk D, Hildebolt C, Smith K, Morwood MJ, Sutikna T, Jatmiko, Saptomo EW, Brunsden B, and Prior F (2006) Response to Comment on "The Brain of LB1, Homo floresiensis". Science 312:999c. Falk D, Hildebolt C, Smith K, Morwood MJ, Sutikna T, Jatmiko, Saptomo EW, Imhof H, Seidler H, and Prior F (2007) Brain shape in human microcephalics and Homo floresiensis. PNAS %R 10.1073/pnas.0609185104:0609185104.
42
Falk D, Redmond JC, Jr., Guyer J, Conroy C, Recheis W, Weber GW, and Seidler H (2000) Early hominid brain evolution: a new look at old endocasts. J Hum Evol 38:695-717. Feldman R (2004) Child Development. Upper Saddle River: Prentice Hall.
Finlay BL, and Darlington RB (1995) Linked regularities in the development and evolution of mammalian brains. Science 268:1578-84. Fox S (1990) Human Physiology. Dubuque: Wm. C. Brown.
Garey LJ (2006) Brodmann's Localisation in the Cerebral Cortex. New York: Springer.
Gibson KR, Rumbaugh D, and Beran M (2001) Bigger is better: primate brain size in relationship to cognition. In KR Gibson and D Falk (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press. Gogtay N, Giedd JN, Lusk L, Hayashi KM, Greenstein D, Vaituzis AC, Nugent TF, 3rd, Herman DH, Clasen LS, Toga AW, Rapoport JL, and Thompson PM (2004) From the Cover: Dynamic mapping of human cortical development during childhood through early adulthood. Proc Natl Acad Sci U S A 101:8174-9. Gur RC, Turetsky BI, Matsui M, Yan M, Bilker W, Hughett P, and Gur RE (1999) Sex differences in brain gray and white matter in healthy young adults: correlations with cognitive performance. J Neurosci 19:4065-72. Gurche J (1982) Early Primate Brain Evolution. In E Armstrong and D Falk (eds.): Primate brain evolution: methods and concepts. New York: Plenum Press. Ho K, Roessmann U, Straumfjord j, and Monroe g (1980) Analysis of Brain weight II Adult brain weight in relation to body height, weight, and surface area. Arch Pathol Lab Med 104:640-645. Hofman M (2001) Brain evolution in hominids: are we at the end of the road. In KR Gibson and D Falk (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press. Holloway RL, Broadfield D, and Yuan M (2004) The Human Fossil Record, Volume 3, Brain Endocasts - The paleoneurological evidence. New York: Willey. 43
Holloway RL, Yuan MS, Broadfield DC, Degusta D, Richards GD, Silvers A, Shapiro JS, and White TD (2002) Missing Omo L338y-6 occipital-marginal sinus drainage pattern: ground sectioning, computer tomography scanning, and the original fossil fail to show it. Anat Rec 266:249-57. Jacob T, Indriati E, Soejono RP, Hsu K, Frayer DW, Eckhardt RB, Kuperavage AJ, Thorne A, and Henneberg M (2006) Pygmoid Australomelanesian Homo sapiens skeletal remains from Liang Bua, Flores: Population affinities and pathological abnormalities 10.1073/pnas.0605563103. PNAS:0605563103.
Kaskan P, and Finlay BL (2001) Encephalization and its developmental structure: how many ways can a brain get big? In KR Gibson and D Falk (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press, pp. 14-29. Le May M (1976) Morphological cerebral asymmetries of modern man, fossil man, and nonhuman primate. Annals of the New York Academy of Sciences 280:349-366. Martin RD, Maclarnon AM, Phillips JL, Dussubieux L, Williams PR, and Dobyns WB (2006) Comment on "The Brain of LB1, Homo floresiensis". Science 312:999; author reply 999. Pakkenberg B, and Gundersen HJ (1997) Neocortical neuron number in humans: effect of sex and age. J Comp Neurol 384:312-20. Passingham R (2006) Cognitive science: brain development and IQ. Nature 440:619-20.
Preuss TM (2001) The discovery of cerebral diversity: an unwelcome scientific revolution. In KR Gibson and D Falk (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press. Purves D, Augustine G, Fitzpatrick D, Katz L, LaMantina A, and McNamara J (1999) Neuroscience. Paris: De Boeck Universite.
44
Radinsky L (1973) Aegyptopithecus endocasts: oldest record of a pongid brain. Am J Phys Anthropol 39:239-47. Radinsky L (1975) Primate brain evolution. Am Sci 63:656-63.
Rakic P, and Kornack D (2001) Neocortical expansion and elaboration during primate evolution: a view from neuroembryology. In KR Gibson and D Falk (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press, pp. 30-56. Semendeferi K (2001) Advances in the study of hominoid brain evolution: magnetic resonance imaging (MRI) and 3-D reconstruction. In D Falk and KR Gibson (eds.): Evolutionary anatomy of the primate cerebral cortex. Cambridge: Cambridge University Press, pp. 257-289.
Semendeferi K, Armstrong E, Schleicher A, Zilles K, and Van Hoesen GW (2001) Prefrontal cortex in humans and apes: a comparative study of area 10. Am J Phys Anthropol 114:224-41. Semendeferi K, and Damasio H (2000) The brain and its main anatomical subdivisions in living hominoids using magnetic resonance imaging. J Hum Evol 38:317-32. Shaw P, Greenstein D, Lerch J, Clasen L, Lenroot R, Gogtay N, Evans A, Rapoport J, and Giedd J (2006) Intellectual ability and cortical development in children and adolescents. Nature 440:676-9. Stephan H, Baron G, and Frahm H (1988) Comparative size of brains and brain components: Comparative primate biology.
Stephan H, Frahm HD, and Baron G (1981) New and revised data on volumes of brain structures in insectivores and primates. Folia Primatol (Basel) 35:1-29. Weber J, Czarnetzki A, and Pusch CM (2005) Comment on "The brain of LB1, Homo floresiensis". Science 310:236; author reply 236. White DD, and Falk D (1999) A quantitative and qualitative reanalysis of the endocast from the juvenile Paranthropus specimen l338y-6 from Omo, Ethiopia. Am J Phys Anthropol 110:399-406.
Zilles K (2005) Evolution of the human brain and comparative cyto- and receptor architecture. In S Dehaene, J Duhamel, M Hauser and G Rizzolatti (eds.): From monkey brain to human brain: A Fyssen Foundation Symposium. Cambridge: MIT Press, pp. 41-56. Zimmer C (2004) Soul made flesh. New York: Free Press.
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FIGURE TITLES AND CREDITS

Figure 1. Brain volume measurements from Stephan et al. (1981) data set. (modified after Rilling, 2006). Box 1 Figure. Scaling: Isometry and Allometry
Figure 2. Double logarithmic scale comparing body weight in grams to brain weight in milligrams of extant primates.
Figure 3. Cranial capacity (a measure of brain size) of fossil hominid specimens derived from Holloway et al. (2004) and Falk et al. (2005). Figure 4. The basic components of a motor neuron.
Figure 5. The human brain in occipital view. Modified from the Comparative Mammalian Brain Collections website http://brainmuseum.org from the University of Wisconsin-Madison Brain Collection. Figure 6. Brodmanns 1909 cytoarchitectonic map of the human brain. Modified after Garey (2006:110). Figure 7. The major subdivisions of the human brain. Figure 8. Major subdivision of the cerebrum.
Figure 9. Primary areas and regions of interest in primate and hominid brain evolution. Figure 10. Human endocast in superior view.
Figure 11. A rubber copy of the human brain and an endocast of the internal cranium of the same individual. Replicas by Bone Clones Inc. Figure 12. Lateral view of a human endocast.
46
Figure 13. The brains of A. Bushbaby B. Macaque C. Chimpanzee drawn to approximately the same size to eliminate the effects of body size. Modified from
the Comparative Mammalian Brain Collections website http://brainmuseum.org from the University of Wisconsin-Madison Brain Collection Figure 14. Percentage of brain volume dedicated to olfactory bulbs, visual cortex, and neocortex from the Stephan et al. (1988) data set. Figure 15. A. Prosimian brain showing the direction of the coronolateral sulcus; B. New World monkey brain showing the central sulcus separating the primary sensory and primary motor areas; C. Old World monkey brain. Figure 16. A. Human brain B. Chimpanzee brain.
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Brain Evolution Watermark

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Brain Evolution Watermark

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Evolution of the Brain:

Neuroanatomy, Development, and Paleontology

HUMAN BRAIN DEVELOPMENT

LATERALIZATION, HANDEDNESS, AND LANGUAGE SEXUAL DIMORPHISM IN THE HUMAN BRAIN

COMPARING NONHUMAN PRIMATE AND HUMAN BRAINS

The indirect or comparative neuroscientific method The direct or paleoneurological method

PRIMATE PALEONEUROLOGY HOMINID PALEONEUROLOGY

Adaptive shifts in primate evolution that impact the brain

GLOSSARY AND KEY WORDS QUESTIONS FOR REVIEW

SUGGESTED FURTHER READING REFERENCES CITED FIGURE TITLES AND CREDITS

Figure 4. The basic components of a motor neuron.

HUMAN BRAIN DEVELOPMENT

LATERALIZATION, HANDEDNESS, AND LANGUAGE

SEXUAL DIMORPHISM IN THE HUMAN BRAIN

COMPARING NONHUMAN PRIMATE AND HUMAN BRAINS

Human Bonobo Chimpanzee Gorilla Orangutan Gibbon

*Left and right sides combined.

Figure 12. Lateral view of a human endocast.

GLOSSARY AND KEY WORDS

Association areas Auditory agnosia Axon

Brain stem Caudal

Central nervous system Chemical synapse Coronolateral sulcus

Electrochemical impulses more commonly known as nerve impulses

Direct method Encephalized Endocasts

Exteroception Frontal pole

Gyri (sing. Gyrus) Gyrification

Indirect method Lateralization

Lesions Lunate sulcus

Neuroglia (glia) Neurons

Occipital lobe Occipital pole Ontogeny

Peripheral nervous system Petalia

Planum temporale Primary areas Proprioception

Prosopagnosia Relative brain size Rostral

Sulci (sing. Sulcus) Superior

QUESTIONS FOR REVIEW

SUGGESTED FURTHER READING

Dunbar R (1998) The social brain hypothesis. Evolutionary Anthropology 6:178109.

FIGURE TITLES AND CREDITS

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