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Saturday, April 18, 2009

Comparative morphology:

The brain of all vertebrates, including humans, consists of three basic divisions: prosencephalon, mesencephalon, and rhombencephalon (Fig. 1). The individual divisions or patterns of the brain do not function separately to bring about a final response; rather, each pattern acts on a common set of connections in the spinal cord.

Lateral views of several vertebrate brains showing evolutionary relationships.
Lateral views of several vertebrate brains showing evolutionary relationships.

Spinal patterns are the final common patterns used by all higher brain pathways to influence all organs of the body. These reflexes are divided into two basic patterns: the monosynaptic arc and the multisynaptic arc. The monosynaptic arc, or myotatic reflex, maintains tonus and posture in vertebrates and consists of two neurons, a sensory and a motor neuron.

The multisynaptic arc, or flexor reflex, is the pattern by which an animal withdraws a part of its body from a noxious stimulus. Both sensory neurons and internuncial neurons send information to brain centers. Coordinated limb movement is based on a connective pattern of neurons at the spinal level.

The structure of the spinal cord and its connections are basically similar among all vertebrates. The major evolutionary changes in the spinal cord have been the increased segregation of cells and fibers of a common function from cells and fibers of other functions and the increase in the length of fibers which connect brain centers with spinal centers. See also Postural equilibrium.

The rhombencephalon of the brain is subdivided into a roof, or cerebellum, and a floor, or medulla oblongata. The medulla is similar to the spinal cord and is divided into a dorsal sensory region and a ventral motor region. It is an integrating and relay area between higher brain centers and the spinal cord. In addition to these nuclei and their connections, the medulla consists of both ascending and descending pathways to and from higher brain centers. The same basic connections occur throughout vertebrates.

In mammals, the cerebellum does not initiate movement; it only times the length of muscle contractions and orders the sequence in which muscles should contract to bring about a movement. The command to initiate a movement is received from the cerebral cortex (Fig. 2). Similarly, the cerebral cortex receives information regarding limb position and state of muscular contraction to ensure that its commands can be carried out by the cerebellum.

Mammalian brain in sagittal section. Cerebellar patterns: tract 1, posterior cerebellar peduncle; 2, middle cerebellar peduncle; 3, anterior cerebellar peduncle.
Mammalian brain in sagittal section. Cerebellar patterns: tract 1, posterior cerebellar peduncle; 2, middle cerebellar peduncle; 3, anterior cerebellar peduncle.

The mesencephalon is divided into a roof or optic tectum and a floor or tegmentum. The tegmentum contains the nuclei of the oculomotor and trochlear cranial nerves and a rostral continuation of the sensory nucleus of the trigeminal cranial nerve.

In the evolution of vertebrates, the prosencephalon develops as two major divisions, the diencephalon and the telencephalon. The diencephalon retains the tubular form and serves as a relay and integrating center for information passing to and from the telencephalon and lower centers. The telencephalon is divided into a pair of cerebral hemispheres and an unpaired telencephalon medium.

There are three divisions of the diencephalon in all vertebrates: an epithalamus which forms the roof of the neural tube, a thalamus which forms the walls of the neural tube, and a hypothalamus which forms the floor of the neural tube. The epithalamus and hypothalamus are primarily concerned with autonomic functions such as homeostasis. The thalamus is subdivided into dorsal and ventral regions. The dorsal region relays and integrates sensory information, and the ventral thalamus relays and integrates motor information. See also Homeostasis; Instinctive behavior.

The telencephalon is the most complex brain division in vertebrates. It is divided into a roof, or pallium, and a floor, or basal region. The pallium is divided into three primary divisions: a medial PI or hippocampal division, a dorsal PII or general pallial division, and a lateral PII division, often called the pyriform pallium.

The most striking change in the telencephalon of land vertebrates involves the PIIIa component. In mammals, it has proliferated with the PIIb component of the dorsal pallium to produce the mammalian neocortex. In all land vertebrates except amphibians, the PIIb and the PIIIa components, along with the corpus striatum (BI and BII), are the highest centers for the analysis of sensory information and motor coordination. The PI, PIIa, PIIIb, BIII, and posterior parts of BI and BII form part of the limbic system which is concerned with behavioral regulation.

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