Brain (2)

Physiology

The functions of the brain depend on the ability of neurons to transmit electrochemical signals to other cells, and their ability to respond appropriately to electrochemical signals received from other cells. The electrical properties of neurons are controlled by a wide variety of biochemical and metabolic processes, most notably the interactions between neurotransmitters and receptors that take place at synapses.[12]

Neurotransmitters and receptors

Neurotransmitters are chemicals that are released at synapses when an action potential activates them—neurotransmitters attach themselves to receptor molecules on the membrane of the synapse's target cell, and thereby alter the electrical or chemical properties of the receptor molecules. With few exceptions, each neuron in the brain releases the same chemical neurotransmitter, or combination of neurotransmitters, at all the synaptic connections it makes with other neurons; this rule is known as Dale's principle.[63] Thus, a neuron can be characterized by the neurotransmitters that it releases. The great majority of psychoactive drugs exert their effects by altering specific neurotransmitter systems. This applies to drugs such as marijuana, nicotine, heroin, cocaine, alcohol, fluoxetine, chlorpromazine, and many others.[64]

The two neurotransmitters that are used most widely in the vertebrate brain are glutamate, which almost always exerts excitatory effects on target neurons, and gamma-aminobutyric acid (GABA), which is almost always inhibitory. Neurons using these transmitters can be found in nearly every part of the brain.[65] Because of their ubiquity, drugs that act on glutamate or GABA tend to have broad and powerful effects. Some general anesthetics act by reducing the effects of glutamate; most tranquilizers exert their sedative effects by enhancing the effects of GABA.[66]

There are dozens of other chemical neurotransmitters that are used in more limited areas of the brain, often areas dedicated to a particular function. Serotonin, for example—the primary target of antidepressant drugs and many dietary aids—comes exclusively from a small brainstem area called the Raphe nuclei.[67] Norepinephrine, which is involved in arousal, comes exclusively from a nearby small area called the locus coeruleus.[68] Other neurotransmitters such as acetylcholine and dopamine have multiple sources in the brain, but are not as ubiquitously distributed as glutamate and GABA.[69]

Electrical Activity

As a side effect of the electrochemical processes used by neurons for signaling, brain tissue generates electric fields when it is active. When large numbers of neurons show synchronized activity, the electric fields that they generate can be large enough to detect outside the skull, using electroencephalography (EEG) [70] or magnetoencephalography (MEG). EEG recordings, along with recordings made from electrodes implanted inside the brains of animals such as rats, show that the brain of a living animal is constantly active, even during sleep.[71] Each part of the brain shows a mixture of rhythmic and nonrhythmic activity, which may vary according to behavioral state. In mammals, the cerebral cortex tends to show large slow delta waves during sleep, faster alpha waves when the animal is awake but inattentive, and chaotic-looking irregular activity when the animal is actively engaged in a task. During an epileptic seizure, the brain's inhibitory control mechanisms fail to function and electrical activity rises to pathological levels, producing EEG traces that show large wave and spike patterns not seen in a healthy brain. Relating these population-level patterns to the computational functions of individual neurons is a major focus of current research in neurophysiology.[71]

Metabolism

All vertebrates have a blood–brain barrier that allows metabolism inside the brain to operate differently from metabolism in other parts of the body. Glial cells play a major role in brain metabolism, by controlling the chemical composition of the fluid that surrounds neurons, including levels of ions and nutrients.[72]

Brain tissue consumes a large amount of energy in proportion to its volume, so large brains place severe metabolic demands on animals. The need to limit body weight in order, for example, to fly, has apparently led to selection for a reduction of brain size in some species, such as bats.[73] Most of the brain's energy consumption goes into sustaining the electric charge (membrane potential) of neurons.[72] Most vertebrate species devote between 2% and 8% of basal metabolism to the brain. In primates, however, the fraction is much higher—in humans it rises to 20–25%.[74] The energy consumption of the brain does not vary greatly over time, but active regions of the cerebral cortex consume somewhat more energy than inactive regions; this forms the basis for the functional brain imaging methods PET, fMRI.[75] and NIRS.[76] The brain typically gets most of its energy from oxygen-dependent metabolism of glucose (i.e., blood sugar),[72] but ketones provide a major alternative source, together with contributions from medium chain fatty acids (octanoic [1] and hexanoic [2] acids), lactate [3], acetate [4], and possibly amino acids.[77]

Functions

From an evolutionary-biological perspective, the function of the brain is to provide coherent control over the actions of an animal. A centralized brain allows groups of muscles to be co-activated in complex patterns; it also allows stimuli impinging on one part of the body to evoke responses in other parts, and it can prevent different parts of the body from acting at cross-purposes to each other.[78]

To generate purposeful and unified action, the brain first brings information from sense organs together at a central location. It then processes this raw data to extract information about the structure of the environment. Next it combines the processed sensory information with information about the current needs of an animal and with memory of past circumstances. Finally, on the basis of the results, it generates motor response patterns that are suited to maximize the welfare of the animal. These signal-processing tasks require intricate interplay between a variety of functional subsystems.[78]

Information processing

The invention of electronic computers in the 1940s, along with the development of mathematical information theory, led to a realization that brains can potentially be understood as information processing systems. This concept formed the basis of the field of cybernetics, and eventually gave rise to the field now known as computational neuroscience.[79] The earliest attempts at cybernetics were somewhat crude in that they treated the brain as essentially a digital computer in disguise, as for example in John von Neumann's 1958 book, The Computer and the Brain.[80] Over the years, though, accumulating information about the electrical responses of brain cells recorded from behaving animals has steadily moved theoretical concepts in the direction of increasing realism.[79]

The essence of the information processing approach is to try to understand brain function in terms of information flow and implementation of algorithms.[79] One of the most influential early contributions was a 1959 paper titled What the frog's eye tells the frog's brain: the paper examined the visual responses of neurons in the retina and optic tectum of frogs, and came to the conclusion that some neurons in the tectum of the frog are wired to combine elementary responses in a way that makes them function as "bug perceivers".[81] A few years later David Hubel and Torsten Wiesel discovered cells in the primary visual cortex of monkeys that become active when sharp edges move across specific points in the field of view—a discovery that eventually brought them a Nobel Prize.[82] Follow-up studies in higher-order visual areas found cells that detect binocular disparity, color, movement, and aspects of shape, with areas located at increasing distances from the primary visual cortex showing increasingly complex responses.[83] Other investigations of brain areas unrelated to vision have revealed cells with a wide variety of response correlates, some related to memory, some to abstract types of cognition such as space.[84]

Theorists have worked to understand these response patterns by constructing mathematical models of neurons and neural networks, which can be simulated using computers.[79] Some useful models are abstract, focusing on the conceptual structure of neural algorithms rather than the details of how they are implemented in the brain; other models attempt to incorporate data about the biophysical properties of real neurons.[85] No model on any level is yet considered to be a fully valid description of brain function, though. The essential difficulty is that sophisticated computation by neural networks requires distributed processing in which hundreds or thousands of neurons work cooperatively—current methods of brain activity recording are only capable of isolating action potentials from a few dozen neurons at a time.[86]

Perception

One of the primary functions of a brain is to extract biologically relevant information from sensory inputs. The human brain is provided with information about light, sound, the chemical composition of the atmosphere, temperature, head orientation, limb position, the chemical composition of the bloodstream, and more. In other animals additional senses may be present, such as the infrared heat-sense of snakes, the magnetic field sense of some birds, or the electric field sense of some types of fish. Moreover, other animals may develop existing sensory systems in new ways, such as the adaptation by bats of the auditory sense into a form of sonar. One way or another, all of these sensory modalities are initially detected by specialized sensors that project signals into the brain.[87]

Each sensory system begins with specialized receptor cells, such as light-receptive neurons in the retina of the eye, vibration-sensitive neurons in the cochlea of the ear, or pressure-sensitive neurons in the skin. The axons of sensory receptor cells travel into the spinal cord or brain, where they transmit their signals to a first-order sensory nucleus dedicated to one specific sensory modality. This primary sensory nucleus sends information to higher-order sensory areas that are dedicated to the same modality. Eventually, via a way-station in the thalamus, the signals are sent to the cerebral cortex, where they are processed to extract biologically relevant features, and integrated with signals coming from other sensory systems.[87]

Motor control

Motor systems are areas of the brain that are directly or indirectly involved in producing body movements, that is, in activating muscles. Except for the muscles that control the eye, which are driven by nuclei in the midbrain, all the voluntary muscles in the body are directly innervated by motor neurons in the spinal cord and hindbrain.[88] Spinal motor neurons are controlled both by neural circuits intrinsic to the spinal cord, and by inputs that descend from the brain. The intrinsic spinal circuits implement many reflex responses, and contain pattern generators for rhythmic movements such as walking or swimming. The descending connections from the brain allow for more sophisticated control.[89]

The brain contains several motor areas that project directly to the spinal cord. At the lowest level are motor areas in the medulla and pons, which control stereotyped movements such as walking, breathing, or swallowing. At a higher level are areas in the midbrain, such as the red nucleus, which is responsible for coordinating movements of the arms and legs. At a higher level yet is the primary motor cortex, a strip of tissue located at the posterior edge of the frontal lobe. The primary motor cortex sends projections to the subcortical motor areas, but also sends a massive projection directly to the spinal cord, through the pyramidal tract. This direct corticospinal projection allows for precise voluntary control of the fine details of movements. Other motor-related brain areas exert secondary effects by projecting to the primary motor areas. Among the most important secondary areas are the premotor cortex, basal ganglia, and cerebellum.[90]

Major areas involved in controlling movement

Area Location Function

• Ventral horn Spinal cord Contains motor neurons that directly activate muscles[91]
• Oculomotor nuclei Midbrain Contains motor neurons that directly activate the eye muscles[92]
• Cerebellum Hindbrain Calibrates precision and timing of movements[46]
• Basal ganglia Forebrain Action selection on the basis of motivation[93]
• Motor cortex Frontal lobe Direct cortical activation of spinal motor circuits
• Premotor cortex Frontal lobe Groups elementary movements into coordinated patterns[94]
• Supplementary motor area Frontal lobe Sequences movements into temporal patterns[95]
• Prefrontal cortex Frontal lobe Planning and other executive functions[96]

In addition to all of the above, the brain and spinal cord contain extensive circuitry to control the autonomic nervous system, which works by secreting hormones and by modulating the "smooth" muscles of the gut.[97] The autonomic nervous system affects heart rate, digestion, respiration rate, salivation, perspiration, urination, and sexual arousal, and several other processes. Most of its functions are not under direct voluntary control.

Arousal

Perhaps the most obvious aspect of the behavior of any animal is the daily cycle between sleeping and waking. Arousal and alertness are also modulated on a finer time scale, though, by an extensive network of brain areas.[98]

A key component of the arousal system is the suprachiasmatic nucleus (SCN), a tiny part of the hypothalamus located directly above the point at which the optic nerves from the two eyes cross. The SCN contains the body's central biological clock. Neurons there show activity levels that rise and fall with a period of about 24 hours, circadian rhythms: these activity fluctuations are driven by rhythmic changes in expression of a set of "clock genes". The SCN continues to keep time even if it is excised from the brain and placed in a dish of warm nutrient solution, but it ordinarily receives input from the optic nerves, through the retinohypothalamic tract (RHT), that allows daily light-dark cycles to calibrate the clock.[99]

The SCN projects to a set of areas in the hypothalamus, brainstem, and midbrain that are involved in implementing sleep-wake cycles. An important component of the system is the reticular formation, a group of neuron-clusters scattered diffusely through the core of the lower brain. Reticular neurons send signals to the thalamus, which in turn sends activity-level-controlling signals to every part of the cortex. Damage to the reticular formation can produce a permanent state of coma.[98]

Sleep involves great changes in brain activity.[100] Until the 1950s it was generally believed that the brain essentially shuts off during sleep,[101] but this is now known to be far from true; activity continues, but patterns become very different. There are two types of sleep: REM sleep (with dreaming) and NREM (non-REM, usually without dreaming) sleep, which repeat in slightly varying patterns throughout a sleep episode. Three broad types of distinct brain activity patterns can be measured: REM, light NREM and deep NREM. During deep NREM sleep, also called slow wave sleep, activity in the cortex takes the form of large synchronized waves, whereas in the waking state it is noisy and desynchronized. Levels of the neurotransmitters norepinephrine and serotonin drop during slow wave sleep, and fall almost to zero during REM sleep; levels of acetylcholine show the reverse pattern.[100]

Homeostasis

For any animal, survival requires maintaining a variety of parameters of bodily state within a limited range of variation: these include temperature, water content, salt concentration in the bloodstream, blood glucose levels, blood oxygen level, and others.[102] The ability of an animal to regulate the internal environment of its body—the milieu intérieur, as pioneering physiologist Claude Bernard called it—is known as homeostasis (Greek for "standing still").[103] Maintaining homeostasis is a crucial function of the brain. The basic principle that underlies homeostasis is negative feedback: any time a parameter diverges from its set-point, sensors generate an error signal that evokes a response that causes the parameter to shift back toward its optimum value.[102] (This principle is widely used in engineering, for example in the control of temperature using a thermostat.)

In vertebrates, the part of the brain that plays the greatest role is the hypothalamus, a small region at the base of the forebrain whose size does not reflect its complexity or the importance of its function.[102] The hypothalamus is a collection of small nuclei, most of which are involved in basic biological functions. Some of these functions relate to arousal or to social interactions such as sexuality, aggression, or maternal behaviors; but many of them relate to homeostasis. Several hypothalamic nuclei receive input from sensors located in the lining of blood vessels, conveying information about temperature, sodium level, glucose level, blood oxygen level, and other parameters. These hypothalamic nuclei send output signals to motor areas that can generate actions to rectify deficiencies. Some of the outputs also go to the pituitary gland, a tiny gland attached to the brain directly underneath the hypothalamus. The pituitary gland secretes hormones into the bloodstream, where they circulate throughout the body and induce changes in cellular activity.[104]

Motivation

According to evolutionary theory, all species are genetically programmed to act as though they have a goal of surviving and propagating offspring. At the level of an individual animal, this overarching goal of genetic fitness translates into a set of specific survival-promoting behaviors, such as seeking food, water, shelter, and a mate.[105] The motivational system in the brain monitors the current state of satisfaction of these goals, and activates behaviors to meet any needs that arise. The motivational system works largely by a reward–punishment mechanism. When a particular behavior is followed by favorable consequences, the reward mechanism in the brain is activated, which induces structural changes inside the brain that cause the same behavior to be repeated later, whenever a similar situation arises. Conversely, when a behavior is followed by unfavorable consequences, the brain's punishment mechanism is activated, inducing structural changes that cause the behavior to be suppressed when similar situations arise in the future.[106]

Every type of animal brain that has been studied uses a reward–punishment mechanism: even worms and insects can alter their behavior to seek food sources or to avoid dangers.[107] In vertebrates, the reward-punishment system is implemented by a specific set of brain structures, at the heart of which lie the basal ganglia, a set of interconnected areas at the base of the forebrain.[50] There is substantial evidence that the basal ganglia are the central site at which decisions are made: the basal ganglia exert a sustained inhibitory control over most of the motor systems in the brain; when this inhibition is released, a motor system is permitted to execute the action it is programmed to carry out. Rewards and punishments function by altering the relationship between the inputs that the basal ganglia receive and the decision-signals that are emitted. The reward mechanism is better understood than the punishment mechanism, because its role in drug abuse has caused it to be studied very intensively. Research has shown that the neurotransmitter dopamine plays a central role: addictive drugs such as cocaine, amphetamine, and nicotine either cause dopamine levels to rise or cause the effects of dopamine inside the brain to be enhanced.[108]

Learning and memory

Almost all animals are capable of modifying their behavior as a result of experience—even the most primitive types of worms. Because behavior is driven by brain activity, changes in behavior must somehow correspond to changes inside the brain. Theorists dating back to Santiago Ramón y Cajal argued that the most plausible explanation is that learning and memory are expressed as changes in the synaptic connections between neurons.[109] Until 1970, however, experimental evidence to support the synaptic plasticity hypothesis was lacking. In 1971 Tim Bliss and Terje Lømo published a paper on a phenomenon now called long-term potentiation: the paper showed clear evidence of activity-induced synaptic changes that lasted for at least several days.[110] Since then technical advances have made these sorts of experiments much easier to carry out, and thousands of studies have been made that have clarified the mechanism of synaptic change, and uncovered other types of activity-driven synaptic change in a variety of brain areas, including the cerebral cortex, hippocampus, basal ganglia, and cerebellum.[111]

Neuroscientists currently distinguish several types of learning and memory that are implemented by the brain in distinct ways:

Working memory is the ability of the brain to maintain a temporary representation of information about the task that an animal is currently engaged in. This sort of dynamic memory is thought to be mediated by the formation of cell assemblies—groups of activated neurons that maintain their activity by constantly stimulating one another.[112]

Episodic memory is the ability to remember the details of specific events. This sort of memory can last for a lifetime. Much evidence implicates the hippocampus in playing a crucial role: people with severe damage to the hippocampus sometimes show amnesia, that is, inability to form new long-lasting episodic memories.[113]

Semantic memory is the ability to learn facts and relationships. This sort of memory is probably stored largely in the cerebral cortex, mediated by changes in connections between cells that represent specific types of information.[114]

Instrumental learning is the ability for rewards and punishments to modify behavior. It is implemented by a network of brain areas centered on the basal ganglia.[115]

Motor learning is the ability to refine patterns of body movement by practicing, or more generally by repetition. A number of brain areas are involved, including the premotor cortex, basal ganglia, and especially the cerebellum, which functions as a large memory bank for microadjustments of the parameters of movement.[116]

Brain (1)

This article is about the brains of all types of animals, including humans. For information specific to the human brain, see Human brain. For other uses, see Brain (disambiguation).

The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few invertebrates such as sponges, jellyfish, adult sea squirts and starfish do not have one, even if diffuse neural tissue is present. It is located in the head, usually close to the primary sensory organs for such senses as vision, hearing, balance, taste, and smell. The brain of a vertebrate is the most complex organ of its body. In a typical human the cerebral cortex (the largest part) is estimated to contain 15–33 billion neurons,[1] each connected by synapses to several thousand other neurons. These neurons communicate with one another by means of long protoplasmic fibers called axons, which carry trains of signal pulses called action potentials to distant parts of the brain or body targeting specific recipient cells.

Physiologically, the function of the brain is to exert centralized control over the other organs of the body. The brain acts on the rest of the body both by generating patterns of muscle activity and by driving secretion of chemicals called hormones. This centralized control allows rapid and coordinated responses to changes in the environment. Some basic types of responsiveness such as reflexes can be mediated by the spinal cord or peripheral ganglia, but sophisticated purposeful control of behavior based on complex sensory input requires the information-integrating capabilities of a centralized brain.

From a philosophical point of view, what makes the brain special in comparison to other organs is that it forms the physical structure that generates the mind. As Hippocrates put it: "Men ought to know that from nothing else but the brain come joys, delights, laughter and sports, and sorrows, griefs, despondency, and lamentations."[2] Through much of history, the mind was thought to be separate from the brain. Even for present-day neuroscience, the mechanisms by which brain activity gives rise to consciousness and thought remain very challenging to understand: despite rapid scientific progress, much about how the brain works remains a mystery. The operations of individual brain cells are now understood in considerable detail, but the way they cooperate in ensembles of millions has been very difficult to decipher. The most promising approaches treat the brain as a biological computer, very different in mechanism from electronic computers, but similar in the sense that it acquires information from the surrounding world, stores it, and processes it in a variety of ways.

This article compares the properties of brains across the entire range of animal species, with the greatest attention to vertebrates. It deals with the human brain insofar as it shares the properties of other brains. The ways in which the human brain differs from other brains are covered in the human brain article. Several topics that might be covered here are instead covered there because much more can be said about them in a human context. The most important is brain disease and the effects of brain damage, covered in the human brain article because the most common diseases of the human brain either do not show up in other species, or else manifest themselves in different ways.

Anatomy

The shape and size of the brains of different species vary greatly, and identifying common features is often difficult.[3] Nevertheless, there are a number of principles of brain architecture that apply across a wide range of species.[4] Some aspects of brain structure are common to almost the entire range of animals species;[5] others distinguish "advanced" brains from more primitive ones, or distinguish vertebrates from invertebrates.[3]

The simplest way to gain information about brain anatomy is by visual inspection, but many more sophisticated techniques have been developed. Brain tissue in its natural state is too soft to work with, but it can be hardened by immersion in alcohol or other fixatives, and then sliced apart for examination of the interior. Visually, the interior of the brain consists of areas of so-called grey matter, with a dark color, separated by areas of white matter, with a lighter color. Further information can be gained by staining slices of brain tissue with a variety of chemicals that bring out areas where specific types of molecules are present in high concentrations. It is also possible to examine the microstructure of brain tissue using a microscope, and to trace the pattern of connections from one brain area to another.[6]

Cellular structure

The brains of all species are composed primarily of two broad classes of cells: neurons and glial cells. Glial cells (also known as glia or neuroglia) come in several types, and perform a number of critical functions, including structural support, metabolic support, insulation, and guidance of development. Neurons, however, are usually considered the most important cells in the brain.[7]

The property that makes neurons unique is their ability to send signals to specific target cells over long distances.[8] They send these signals by means of an axon, which is a thin protoplasmic fiber that extends from the cell body and projects, usually with numerous branches, to other areas, sometimes nearby, sometimes in distant parts of the brain or body. The length of an axon can be extraordinary: for example, if a pyramidal cell of the cerebral cortex were magnified so that its cell body became the size of a human body, its axon, equally magnified, would become a cable a few centimeters in diameter, extending more than a kilometer.[9] These axons transmit signals in the form of electrochemical pulses called action potentials, which last less than a thousandth of a second and travel along the axon at speeds of 1–100 meters per second. Some neurons emit action potentials constantly, at rates of 10–100 per second, usually in irregular patterns; other neurons are quiet most of the time, but occasionally emit a burst of action potentials.[10]

Axons transmit signals to other neurons by means of specialized junctions called synapses. A single axon may make as many as several thousand synaptic connections with other cells.[11] When an action potential, traveling along an axon, arrives at a synapse, it causes a chemical called a neurotransmitter to be released. The neurotransmitter binds to receptor molecules in the membrane of the target cell.[12]

Synapses are the key functional elements of the brain.[13] The essential function of the brain is cell-to-cell communication, and synapses are the points at which communication occurs. The human brain has been estimated to contain approximately 100 trillion synapses;[14] even the brain of a fruit fly contains several million.[15] The functions of these synapses are very diverse: some are excitatory (excite the target cell); others are inhibitory; others work by activating second messenger systems that change the internal chemistry of their target cells in complex ways.[13] A large fraction of synapses are dynamically modifiable; that is, they are capable of changing strength in a way that is controlled by the patterns of signals that pass through them. It is widely believed that activity-dependent modification of synapses is the brain's primary mechanism for learning and memory.[13]

Most of the space in the brain is taken up by axons, which are often bundled together in what are called nerve fiber tracts. Many axons are wrapped in thick sheaths of a fatty substance called myelin, which serves to greatly increase the speed of signal propagation. Myelin is white, so parts of the brain filled exclusively with nerve fibers appear as light-colored white matter, in contrast to the darker-colored grey matter that marks areas with high densities of neuron cell bodies.[16]

Evolution

The generic bilaterian nervous system

Except for a few primitive types such as sponges (which have no nervous system[17]) and cnidarians (which have a nervous system consisting of a diffuse nerve net[17]), all living multicellular animals are bilaterians, meaning animals with a bilaterally symmetric body shape (that is, left and right sides that are approximate mirror images of each other).[18] All bilaterians are thought to have descended from a common ancestor that appeared early in the Cambrian period, 550–600 million years ago, and it has been hypothesized that this common ancestor had the shape of a simple tubeworm with a segmented body.[18] At a schematic level, that basic worm-shape continues to be reflected in the body and nervous system architecture of all modern bilaterians, including vertebrates.[19] The fundamental bilateral body form is a tube with a hollow gut cavity running from the mouth to the anus, and a nerve cord with an enlargement (a ganglion) for each body segment, with an especially large ganglion at the front, called the brain. The brain is small and simple in some species, such as nematode worms; in other species, including vertebrates, it is the most complex organ in the body.[3] Some types of worms, such as leeches, also have an enlarged ganglion at the back end of the nerve cord, known as a "tail brain".[20]

There are a few types of existing bilaterians that lack a recognizable brain, including echinoderms, tunicates, and a group of primitive flatworms called Acoelomorpha. It has not been definitively established whether the existence of these brainless species indicates that the earliest bilaterians lacked a brain, or whether their ancestors evolved in a way that led to the disappearance of a previously existing brain structure.[21]

Invertebrates

This category includes arthropods, molluscs, and numerous types of worms. The diversity of invertebrate body plans is matched by an equal diversity in brain structures.[22]

Two groups of invertebrates have notably complex brains: arthropods (insects, crustaceans, arachnids, and others), and cephalopods (octopuses, squids, and similar molluscs).[23] The brains of arthropods and cephalopods arise from twin parallel nerve cords that extend through the body of the animal. Arthropods have a central brain with three divisions and large optical lobes behind each eye for visual processing.[23] Cephalopods such as the octopus and squid have the largest brains of any  invertebrates.[24]

There are several invertebrate species whose brains have been studied intensively because they have properties that make them convenient for experimental work:

Fruit flies (Drosophila), because of the large array of techniques available for studying their genetics, have been a natural subject for studying the role of genes in brain development.[25] In spite of the large evolutionary distance between insects and mammals, many aspects of Drosophila neurogenetics have turned out to be relevant to humans. The first biological clock genes, for example, were identified by examining Drosophila mutants that showed disrupted daily activity cycles.[26] A search in the genomes of vertebrates turned up a set of analogous genes, which were found to play similar roles in the mouse biological clock—and therefore almost certainly in the human biological clock as well.[27]

The nematode worm Caenorhabditis elegans, like Drosophila, has been studied largely because of its importance in genetics.[28] In the early 1970s, Sydney Brenner chose it as a model system for studying the way that genes control development. One of the advantages of working with this worm is that the body plan is very stereotyped: the nervous system of the hermaphrodite morph contains exactly 302 neurons, always in the same places, making identical synaptic connections in every worm.[29] Brenner's team sliced worms into thousands of ultrathin sections and photographed every section under an electron microscope, then visually matched fibers from section to section, to map out every neuron and synapse in the entire body.[30] Nothing approaching this level of detail is available for any other organism, and the information has been used to enable a multitude of studies that would not have been possible without it.[31]

The sea slug Aplysia was chosen by Nobel Prize-winning neurophysiologist Eric Kandel as a model for studying the cellular basis of learning and memory, because of the simplicity and accessibility of its nervous system, and it has been examined in hundreds of experiments.[32]

Vertebrates

The first vertebrates appeared over 500 million years ago (Mya), during the Cambrian period, and may have resembled the modern hagfish in form.[33] Sharks appeared about 450 Mya, amphibians about 400 Mya, reptiles about 350 Mya, and mammals about 200 Mya. No modern species should be described as more "primitive" than others, strictly speaking, since each has an equally long evolutionary history—but the brains of modern hagfishes, lampreys, sharks, amphibians, reptiles, and mammals show a gradient of size and complexity that roughly follows the evolutionary sequence. All of these brains contain the same set of basic anatomical components, but many are rudimentary in the hagfish, whereas in mammals the foremost part (the telencephalon) is greatly elaborated and expanded.[34]

Brains are most simply compared in terms of their size. The relationship between brain size, body size and other variables has been studied across a wide range of vertebrate species. As a rule, brain size increases with body size, but not in a simple linear proportion. In general, smaller animals tend to have larger brains, measured as a fraction of body size: the animal with the largest brain-size-to-body-size ratio is the hummingbird. For mammals, the relationship between brain volume and body mass essentially follows a power law with an exponent of about 0.75.[35] This formula describes the central tendency, but every family of mammals departs from it to some degree, in a way that reflects in part the complexity of their behavior. For example, primates have brains 5 to 10 times larger than the formula predicts. Predators tend to have larger brains than their prey, relative to body size.[36]

All vertebrate brains share a common underlying form, which appears most clearly during early stages of embryonic development. In its earliest form, the brain appears as three swellings at the front end of the neural tube; these swellings eventually become the forebrain, midbrain, and hindbrain (the prosencephalon, mesencephalon, and rhombencephalon, respectively). At the earliest stages of brain development, the three areas are roughly equal in size. In many classes of vertebrates, such as fish and amphibians, the three parts remain similar in size in the adult, but in mammals the forebrain becomes much larger than the other parts, and the midbrain becomes very small.[37]

The brains of vertebrates are made of very soft tissue.[38] Living brain tissue is pinkish on the outside and mostly white on the inside, with subtle variations in color. Vertebrate brains are surrounded by a system of connective tissue membranes called meninges that separate the skull from the brain. Blood vessels enter the central nervous system through holes in the meningeal layers. The cells in the blood vessel walls are joined tightly to one another, forming the so-called blood–brain barrier, which protects the brain from toxins that might enter through the bloodstream.[39]

Neuroanatomists usually divide the vertebrate brain into six main regions: the telencephalon (cerebral hemispheres), diencephalon (thalamus and hypothalamus), mesencephalon (midbrain), cerebellum, pons, and medulla oblongata. Each of these areas has a complex internal structure. Some parts, such as the cerebral cortex and cerebellum, consist of layers that are folded or convoluted to fit within the available space. Other parts, such as the thalamus and hypothalamus, consist of clusters of many small nuclei. Thousands of distinguishable areas can be identified within the vertebrate brain based on fine distinctions of neural structure, chemistry, and connectivity.[38]

Although the same basic components are present in all vertebrate brains, some branches of vertebrate evolution have led to substantial distortions of brain geometry, especially in the forebrain area. The brain of a shark shows the basic components in a straightforward way, but in teleost fishes (the great majority of existing fish species), the forebrain has become "everted", like a sock turned inside out. In birds, there are also major changes in forebrain structure.[40] These distortions can make it difficult to match brain components from one species with those of another species.[41]

Here is a list of some of the most important vertebrate brain components, along with a brief description of their functions as currently understood:

The medulla, along with the spinal cord, contains many small nuclei involved in a wide variety of sensory and motor functions.[42]

The pons lies in the brainstem directly above the medulla. Among other things, it contains nuclei that control sleep, respiration, swallowing, bladder function, equilibrium, eye movement, facial expressions, and posture.[43]

The hypothalamus is a small region at the base of the forebrain, whose complexity and importance belies its size. It is composed of numerous small nuclei, each with distinct connections and neurochemistry. The hypothalamus regulates sleep and wake cycles, eating and drinking, hormone release, and many other critical biological functions.[44]

The thalamus is another collection of nuclei with diverse functions. Some are involved in relaying information to and from the cerebral hemispheres. Others are involved in motivation. The subthalamic area (zona incerta) seems to contain action-generating systems for several types of "consummatory" behaviors, including eating, drinking, defecation, and copulation.[45]

The cerebellum modulates the outputs of other brain systems to make them precise. Removal of the cerebellum does not prevent an animal from doing anything in particular, but it makes actions hesitant and clumsy. This precision is not built-in, but learned by trial and error. Learning how to ride a bicycle is an example of a type of neural plasticity that may take place largely within the cerebellum.[46]

The optic tectum allows actions to be directed toward points in space, most commonly in response to visual input. In mammals it is usually referred to as the superior colliculus, and its best-studied function is to direct eye movements. It also directs reaching movements and other object-directed actions. It receives strong visual inputs, but also inputs from other senses that are useful in directing actions, such as auditory input in owls and input from the thermosensitive pit organs in snakes. In some fishes, such as lampreys, this region is the largest part of the brain.[47] The superior colliculus is part of the midbrain.

The pallium is a layer of gray matter that lies on the surface of the forebrain. In reptiles and mammals, it is called the cerebral cortex. Multiple functions involve the pallium, including olfaction and spatial memory. In mammals, where it becomes so large as to dominate the brain, it takes over functions from many other brain areas. In many mammals, the cerebral cortex consists of folded bulges called gyri that create deep furrows or fissures called sulci. The folds increase the surface area of the cortex and therefore increase the amount of gray matter and the amount of information that can be processed.[48]

The hippocampus, strictly speaking, is found only in mammals. However, the area it derives from, the medial pallium, has counterparts in all vertebrates. There is evidence that this part of the brain is involved in spatial memory and navigation in fishes, birds, reptiles, and mammals.[49]

The basal ganglia are a group of interconnected structures in the forebrain. The primary function of the basal ganglia appears to be action selection: they send inhibitory signals to all parts of the brain that can generate motor behaviors, and in the right circumstances can release the inhibition, so that the action-generating systems are able to execute their actions. Reward and punishment exert their most important neural effects by altering connections within the basal ganglia.[50]

The olfactory bulb is a special structure that processes olfactory sensory signals and sends its output to the olfactory part of the pallium. It is a major brain component in many vertebrates, but is greatly reduced in primates.[51]

Mammals

The most obvious difference between the brains of mammals and other vertebrates is in terms of size. On average, a mammal has a brain roughly twice as large as that of a bird of the same body size, and ten times as large as that of a reptile of the same body size.[52]

Size, however, is not the only difference: there are also substantial differences in shape. The hindbrain and midbrain of mammals are generally similar to those of other vertebrates, but dramatic differences appear in the forebrain, which is greatly enlarged and also altered in structure.[53] The cerebral cortex is the part of the brain that most strongly distinguishes mammals. In non-mammalian vertebrates, the surface of the cerebrum is lined with a comparatively simple three-layered structure called the pallium. In mammals, the pallium evolves into a complex six-layered structure called neocortex or isocortex.[54] Several areas at the edge of the neocortex, including the hippocampus and amygdala, are also much more extensively developed in mammals than in other vertebrates.[53]

The elaboration of the cerebral cortex carries with it changes to other brain areas. The superior colliculus, which plays a major role in visual control of behavior in most vertebrates, shrinks to a small size in mammals, and many of its functions are taken over by visual areas of the cerebral cortex.[52] The cerebellum of mammals contains a large portion (the neocerebellum) dedicated to supporting the cerebral cortex, which has no counterpart in other vertebrates.[55]

Primates

The brains of humans and other primates contain the same structures as the brains of other mammals, but are generally larger in proportion to body size.[59] The most widely accepted way of comparing brain sizes across species is the so-called encephalization quotient (EQ), which takes into account the nonlinearity of the brain-to-body relationship.[56] Humans have an average EQ in the 7-to-8 range, while most other primates have an EQ in the 2-to-3 range. Dolphins have values higher than those of primates other than humans,[57] but nearly all other mammals have EQ values that are substantially lower.

Most of the enlargement of the primate brain comes from a massive expansion of the cerebral cortex, especially the prefrontal cortex and the parts of the cortex involved in vision.[60] The visual processing network of primates includes at least 30 distinguishable brain areas, with a complex web of interconnections. It has been estimated that visual processing areas occupy more than half of the total surface of the primate neocortex.[61] The prefrontal cortex carries out functions that include planning, working memory, motivation, attention, and executive control. It takes up a much larger proportion of the brain for primates than for other species, and an especially large fraction of the human brain.[62]

The Missing Link

Every few years scientists unearth the bones of humanity's forefathers. From Lucy to the Hobbits of Flores Island -- we are gradually seeing building the puzzle of mankind's evolution.

1. Neanderthal Stocky and squat and well suited for the cold, Neanderthals looked distinctly different from modern humans. But they were like us in other ways: they buried their dead, cared for their sick and injured and may have been capable of language and music. Scientists recently put together a complete Neanderthal skeleton and are working on the genome.
2. Cro-Magnon These people looked identical to modern humans and lived in Europe between 35,000 and 10,000 years ago. Their cave paintings and sculptures are the earliest known examples of art by a prehistoric people.
3. Homo floresiensis It turns out those Floresians were actually on to something. For centuries, their mythology described a race of very small human-like creatures called the Ebu Gogo. Hardly anyone took them seriously, however, until 2003, when word broke that a new species of diminutive hominids was discovered on the Indonesian island
4. Homo erectus For H. erectus, it may have paid to be dense. According to one theory, males rammed each other with their thick skulls in order to win females. H. erectus is generally believed to be the direct ancestor of modern humans and also the first hominid to live in caves and tame fire.
5. Homo ergaster Scientists can't decide whether this African hominid is just a failed predecessor of H. erectus or the rightful ancestor of modern humans. It had a thinner skull than H. erectus and was more proficient at making tools and using fire.
6. Homo habilis Many scientists believe H. habilis is the missing link between the ape-like hominids like Lucy and the more human-like ones that came after. It had long ape-like arms but walked on two feet and was capable of creating crude tools.
7. Paranthropus boisei If P. boisei and its relatives weren’t such picky eaters, we might not be here to wonder about them. They split from the line leading to modern human some 2 million years ago and lived alongside our ancestors for millions of years, but died out after failing to adapt their diets
8. Paranthropus aethiopicus This early ape-like hominid walked on two legs and lived between 2.8 million and 2.2 million years ago. Based on skull measurements, scientists concluded this species had the smallest adult hominid brain ever discovered.
9. Australopithecus africanus A. africanus was an early descendent of Lucy and lived in Southern Africa between 2 million and 3 million years ago. Its brain was larger than Lucy's and its facial features were more human-like.
10. Australopithecus afarensis The most famous member of this species is Lucy, an adult female skeleton discovered in 1974 and nicknamed after a Beatles song. Lucy lived about 3.18 million years ago and was fully capable of walking and running on two legs.

8 Steps to plans your Succesful future

Here is several steps to plan your future...

1. Go to a quiet place , where you can think without being interrupted.
2. Go through your life thoroughly. What is your gift? What do you want to be in the future? What is your passion? (i.e., something you
   
   would love to do even if you were not paid to do it.)
3. Write it down. Most of the people who have achieved their dreams took the first step towards achieving them: they wrote them down.
4. Plan a route to get there. Write down what ever it will take you to reach your dream.
5. Put your piece of paper where you can see it and remind yourself of your goals.
6. Work your plan. Put it into action by following the route to your dream. It may mean adjusting a lot of things in your life, but it will be worth it one day.
7. Start a savings account and deposit 25% of your monthly income. Then when you need to buy a car, etc., you'll already have a lot of what you'll need.
8. Dismiss negativity and begin to speak your future in the present. This will assimilate into your mind and help you to persevere towards your dream.

tips

• Talk to your parents, guidance counselors, etc., about what they think you would be good at and how to get there. You may not want to, but remember, they went through this too.
• The difference between a big shot and a little shot is that a big shot is just a little shot who kept on shooting!