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Zool448 7 January 2003 Bosma Lab Early motor neuron development The hindbrain is the site of the cranial nerves, some of which (trigeminal, facial) include motor neurons innervating a unique type of muscle (different origin than other body muscles). Bosma Lab How do we identify the neurons? We inject the targets of the neurons with a red dextran dye, and wait for the dye to get transported up to the somas, in the CNS. Then we use an intracellular Ca2+-indicating dye to examine the physiology of these developing neurons. open dextran 1 Zool448 7 January 2003 Bosma Lab We are sure we have the right cells We can fix the tissue, and look at the dextran signal (red), and counter-stain with an antibody that is specific to motor neurons (green). Bosma Lab The activity changes over time Activity is recorded from individual cells, and each cell’s activity is plotted in a different color on these graphs. At E11.0, the cells are active, but each is independent. Twelve hours later, at E11.5, all the cells are active synchronously. 2 Zool448 7 January 2003 Zebra Finch Female Male 3 Zool448 7 January 2003 Sample zebra finch song kHz 8 0 500 ms The Song System (simplified) HVc Song Production Auditory input LMAN RA Learning X DLM VTA s IIt nX Respiration Syrinx (vocal organ) 4 Zool448 7 January 2003 HVc L-MAN RA Area X DLM Luo et al., 2001 Luo et al., 2001 5 Zool448 7 January 2003 Basic properties of neurons Neurons are a specialized cell type Neurons have the components common to most cells: they have a nucleus which contains DNA, and where genes are transcribed; they have a cytoplasm in which most translation and protein synthesis begins; they have a plasma membrane that surrounds the entire cells; they have an elaborate cytoskeleton to support the cell and carry out transport functions. Basic properties of neurons Neurons are unique in several ways They have a cell body (called the soma), where the nucleus is. They have cellular processes, called dendrites and axons. These processes can be very elaborate, and the axons may be very long. The general function of dendrites is to receive information from other neurons. Dendrites Soma Axon The general function of axons is to transmit information to other cells, such as neurons or muscles. 6 Zool448 7 January 2003 Basic properties of neurons Types of neurons We will concentrate on three different types of neurons: 1. Sensory neurons, that receive information from the outside world. This will include tactile and visual information. 2. Motor neurons, that cause muscle contraction in the periphery of the body. 3. Interneurons, which receive (and often modify) information from neurons, then pass it on to other neurons. Basic properties of neurons Types of neurons 7 Zool448 7 January 2003 Basic properties of neurons Neurons are organized into groups Neurons are usually localized into groups of cell bodies, which underlie the functions of the nervous system. The nervous system is divided into the central nervous system (CNS; brain and spinal cord), and the peripheral nervous system (PNS). In the CNS, a group of cells is usually called a nucleus. Different parts of the brain contain specific nuclei; each nucleus is related to specific modalities (senses) or functions. Groups of cells located in the PNS are usually called ganglia; these would include the sympathetic ganglia, which perform autonomic functions, and sensory ganglia. The PNS also includes the peripheral nerves. Neurons make up nervous systems Invertebrates Hydra – nerve net Starfish – radial nervous system (symmetric) with ganglia Leech – bilaterally symmetric nervous system with ganglia Grasshopper – bilaterally symmetric with ganglia gathered towards the head 8 Zool448 7 January 2003 Neurons make up nervous systems Vertebrates have a dorsal CNS Animals classified as higher on the evolutionary tree have a tendency towards cephalization, a larger and more prominent head. A major change as animals evolved is an increase in the convolutions and complexity of the cortex. The axes of the CNS Rostral-caudal, anterior-posterior Rostral is defined as being towards the nose, while caudal is defined as being towards the tail. Dorsal means towards the back, ventral means towards the front. In lower animals, the axis of the brain is the same as the body axis. Rostral = nose Caudal = tail Dorsal = back Ventral = front 9 Zool448 7 January 2003 The axes of the CNS Rostral-caudal, anterior-posterior In higher animals, the axis of the rostral CNS bends, and is curved almost 110o from the spinal cord axis: thus, by convention, the rostral CNS axis nomenclature differs from that in the spinal cord. Rostral = nose Caudal = back of head Dorsal = top of skull Ventral = jaw The axes of the CNS Medial-lateral Medial means towards the midline of the brain or animal, while lateral is to the side. 10 Zool448 7 January 2003 The planes of section The major types of brain sections are shown: Horizontal sections move from the top of the head to the bottom. Coronal sections move from the front of the head to the back (seen as if wearing a crown). Sagittal sections begin in the midline of the brain and give a side view of the brain; parasagittal sections are off-midline. Divisions of the CNS Spinal cord – most caudal The spinal cord receives sensory information from the trunk and limbs of the body, and controls the motor neurons to that portion of the body. It is divided into four major segments, and contains the 31 pairs of spinal nerves, each of which carries the sensory information from a particular area as well as the motor output for that area. Spinal cord 11 Zool448 7 January 2003 Divisions of the CNS The brain stem contains three parts The brain stem is made up of the medulla, pons and midbrain. It contains the cranial nerves (except olfactory), receiving sensation from the face and special senses. They control the motor components of the face and neck. There are also projection neurons that extend throughout the brain from the reticular formation that control awareness. Brain stem Midbrain Pons Medulla Divisions of the CNS The cerebellum coordinates motor skills The cerebellum receives vestibular information, posture and sensory information from the body, and coordinates that information with visual inputs to allow the fine tuning of posture and body position, and the learning of motor tasks. Cerebellum 12 Zool448 7 January 2003 Divisions of the CNS The diencephalon contains two parts The diencephalon contains two major parts. The thalamus is a major relay center for visual, auditory and vestibular information to the cortex. Diencephalon The hypothalamus controls reproduction and endocrine systems as well as the circadian clock and general body homeostasis. Thalamus Hypothalamus Divisions of the CNS The cerebral hemispheres The cerebral hemispheres are made up of the cerebral cortex on the surface of the brain, which is involved in cognition and planning of motor activities. The three deeper structures are the basal ganglia controlling fine movement, the amygdala controlling emotion and behavior, while the hippocampus is concerned with memory. Basal ganglia Cerebral hemispheres 13 Zool448 7 January 2003 Divisions of the CNS The cerebral hemispheres The cerebral cortex is divided into lobes: the frontal lobe plans actions, the parietal lobe is concerned with representation of the body, the occipital lobe is where vision is mediated, and the temporal lobe is concerned with audition and some aspects of learning. Neurons mediate brain functions Neurons signal extremely rapidly Because of neuron signaling speed, we can act and move quickly. The mechanism of rapid signaling is via electrical impulses called action potentials, which are caused by the opening and closing of ion channel proteins localized in the plasma membrane. Neurons convert this electrical signal into a chemical signal at the synapse, where information is passed to the next cell. The receiving cell (the post-synaptic cell) has specialized receptor proteins for responding to the chemical signal from the first cell cell (pre-synaptic cell). Via this signaling, the post-synaptic cell can then fire action potentials. 14 Zool448 7 January 2003 When would this be used? The knee-jerk response This reflex includes four neurons, two (opposing) muscles and a stimulus (hammer). The sensory neuron stretch receptor in the muscle transduces the input (feels the thwack of the hammer), and signals to two neurons. When would this be used? The knee-jerk response – part 1 The motor neuron causes the quadriceps to contract. 15 Zool448 7 January 2003 When would this be used? The knee-jerk response The motor neuron causes the quadriceps to contract. When would this be used? The knee-jerk response – part 2 The motor neuron causes the quadriceps to contract. An interneuron inverts the signal, and causes the opposing muscle to relax. Figure 6-6 16 Zool448 7 January 2003 Information is passed along What is that motor neuron experiencing? The sensory neuron sends a chemical signal at the synapse (in the form of neurotransmitter) to the motor neuron. This is converted to an electrical signal that is sent to the muscle, through another synapse. Figure 6-1 Neurons are complex Motor neurons are large and complex cells, with many dendritic branches to receive the inputs from other cells, and axons to send the information from the motor neuron. Figure 1-3 17 Zool448 7 January 2003 Example: sensory neuron Signaling is directional and proportional to the stimulus Example: sensory neuron Signaling is directional and proportional to the stimulus 18