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Unit 2: What are the building Overview blocks of our brains? In the last unit we discovered that complex brain functions occur as individual structures in the brain work together like an orchestra. We also discussed one of the limitations of our new visualization techniques – that they only sample populations of hundreds or thousands of neurons, so they don’t give us any information about how the individual cells of the nervous system work together. So now we’re going to take another step back and dial down our focus to the primary building blocks of our brains, the neurons and the glial cells. In this unit we will explore how these basic cells are built and how they work, and importantly what can go wrong when these building blocks are diseased and their functions are compromised. Remember our graphic from the beginning of this workbook? This unit focuses on the neuron, which is the building block of our brains. LESSON 2.1 WORKBOOK What is the structure of a neuron? DEFINITIONS OF TERMS Neuron – cells of the nervous system that are specialized for the reception, conduction and transmission of electrochemical signals. This unit introduces you to the building blocks of our brains: neurons and glia cells. In this lesson, we will begin our exploration of how the brain is put together by investigating why neurons have such complex structures and how these structures allow the neurons to perform highly specialized functions What are neurons? For a complete list of defined terms, see the Glossary. Wo r k b o o k Lesson 2.1 Neurons are the most important functional cells in our nervous system. The adult human brain contains roughly 86 billion individual neurons. Each neuron is interconnected, forming a precise network. Within that network neurons are assembled into many different kinds of functionally distinct regions (like Broca’s area for example). As we saw in the last lesson these regions interact with each other to produce our perception of the external world, to fix our attention on the responses that need to be made, and to control our bodily functions. Our first step in understanding the brain, therefore, has to be to understand the neuron – how it is put together and how it works. Neurons are cells with highly complex structures, much more complex than any other cell in the body. Wiggle your big toe. The neuron that controls that wiggle starts off in the spinal cord somewhere in your upper chest and ends up at your big toe, a distance that would be tens of meters if you were a giraffe (which don’t have toes, but whatever, you get the point). Neurons are different from other cells in a number of ways especially because, unlike most cells, neurons don’t divide — the number of neurons you had when you are born is the maximum you will ever have. This means that when a neuron is damaged the only possibility you have to restore its function is to fix it, you can’t simply make another one to take its place, like you could in the liver. In the peripheral nervous system you can fix damaged neurons so that they’ll grow slowly back to make their original connections. The central nervous system is different. When a CNS neuron is damaged it cannot regrow long distances to repair its connections. Why? No one really knows. (Interestingly, CNS neurons can grow in lower vertebrates like fish). One potential reason why our CNS neurons aren’t able to regrow lays in the hypothesis that all of our complex behaviors demand a neuronal network with a very precise architecture. Meaning that, CNS neurons have had to trade off the ability to regrow, so that the network remains stable. Even so, some nervous system damage can be repaired if we can induce neurons to rewire over short distances. Can neurons in the PNS repair themselves? What would this mean in regards to recovery after an injury to the PNS? __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ Can neurons in the CNS repair themselves? What would this mean in regards to recovery after an injury to the CNS? __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ __________________________________ 38 LESSON READING What are the three functional regions of the neuron? Neurons have three distinct functional regions The typical neuron contains three different regions, each of which looks different and each of which has its own specialized function (Figure 1). These regions are: DEFINITIONS OF TERMS Cell body – part of the neuron containing the nucleus, but not including the axon and dendrites. Also called the soma. Endoplasmic reticulum – organelle in the cell that forms a network of tubules and vesicles. It functions to synthesize proteins and lipids as well as metabolize carbohydrates. Nucleus – the DNA containing structures of cells. For a complete list of defined terms, see the Glossary. Wo r k b o o k Lesson 2.1 • • • The cell body The dendrites The axon Dendrites Axon Cell Body Ini2al Segment Synapse Presynap2c cell Postsynap2c cell Figure 1: Neuron structure. Neurons have three distinct regions: the dendrites, the cell body, and the axon. The cell body The cell body (also sometimes called the soma) is the metabolic center of the neuron (Figure 2). It contains the nucleus, which stores the genes of the cell in chromosomes, and the smooth and rough endoplasmic reticulum, which are the sites where proteins are synthesized. It also contains the lysosomes that degrade proteins that have become old or damaged. Because the ribosomes are mostly concentrated in the cell body, protein synthesis primarily occurs there and in the dendrites that are closest to it. Because of this, a major role of the cell body is to package the proteins it has made so they can be transported over long distances down the leg and into the foot to our big toe (or our little finger etc.). Similarly, because the cell body is also the site were lysosomes are concentrated, any big toe protein that has reached its sell-by date needs to be transported back up the leg to the cell body for destruction. Keeping all the parts of the neuron supplied with protein is a major task carried out by the cell body. Figure 2: Cell body. The cell body is the metabolic center of the cell and contains all the cellular organelles required to support cell life: the nucleus, mitochondria, ribosomes, rough and smooth endoplasmic reticulum. ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ Name two important functions carried out by the neuron’s cell body. ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ __________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ 39 LESSON READING We can identify two types of outgrowths sprouting off from the cell body, the dendrites and the axon. The dendrites DEFINITIONS OF TERMS Dendrites — branched projection(s) of a neuron that functions as the receptive area of a neuron. Dendritic spines — tiny spikes of various shapes that are located on the surfaces of many dendrites and are the sites of synapses. For a complete list of defined terms, see the Glossary. Wo r k b o o k Lesson 2.1 Most neurons have several dendrites (Figure 3). These dendrites branch out from the cell body in a shape that makes them look like a tree. In fact the dendrites are often called ‘the dendritic tree’. The dendritic tree is the main region of the neuron that receives signals. These signals can come in the form of sensations from the environment. Alternatively, in the depths of the neuronal network they may come from other neurons. The role of the Figure 3: Dendrites. The dendritic arbor of two dendrites is to convert these signals, which may neurons (a Purkinje neuron on the left, and a senbe in the form of physical signals if they are from sory neuron on the righ) illustrating the extensive the environment (such as light, sound or touch) or branching of dendrites.. chemicals if they are from other neurons, into an electrical signal. Dendrites do this by changing the electrical properties of their membranes via depolarization or hyperpolarization. We will talk more about the important processes of depolarization and hyperpolarization later on in this unit. Each of our sensory systems contains unique neurons that are specialized to detect specific types of sensory stimuli in the environment. The dendrites from these neurons are able to convert these stimuli into a neural response that the brain can understand. For example, different types of sensory dendrites in our skin are uniquely tuned to detect changes in pressure. They then convert the physical sensation of pressure into a neural response by depolarizing or hyperpolarizing their membranes. The branches of the dendritic tree often have many hundreds of thousands of little twigs that we call dendritic spines because they look like spikes (Figure 4). Each Figure 4: Dendritic spines. Dendrites have small protuberances called spines. dendritic spine usually contains one synapse, which is an Each spine can contain a synapse. exact area where the dendrite can receive a signal, whether from the environment or from another neuron. You can appreciate that if a single dendritic tree has hundreds of thousands of spines, then it can have hundreds of thousands of different inputs. Remember that there are also 86 billion neurons — makes you appreciate that trying to understand how everything is connected is a massive task. No wonder neuroscientists were excited by the development of supercomputers! What is the function of the dendritic tree? ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ Which kind of neuron has more inputs: a neuron without dendritic spines, or a neuron with dendiritc spines? Why? ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ __________________________________ 40 LESSON READING What is the function of the axon? The axon DEFINITIONS OF TERMS Action potential – the electrical signal of the axon. Axon – projection of a neuron that functions to conduct electrical impulses away from a neuron’s cell body. Presynaptic cell – neuron located before the synapse, and thus sending the signal Postsynaptic cell – neuron located after the synapse, and thus receiving the signal For a complete list of defined terms, see the Glossary. Wo r k b o o k Lesson 2.1 The other type of sprout we can detect coming off the cell body is the axon. Unlike the branches of the dendritic tree, which are tapered just like real branches, the axon can be identified because it looks just like a cylindrical tube. There is usually only one axon per neuron. The axon grows out from a specialized region of the cell body called the axon hillock or initial segment. This structure is important because the axon is the main transmitting or conducting unit of the neuron, conveying electrical signals from the dendritic tree down to its very tip. In our big toe analogy, the axon would convey the signal from dendrites in the spinal cord along your leg to tell your muscles to wiggle your toe. The axon hillock gathers together all the signals the neuron has received from the dendritic tree, converts them into the single output response and sends them down the axon. This output response is an electrical signal called the action potential. We will focus on how the action potential is made and transported in another lesson in this unit. Many axons split into several branches at their tips (like the roots of the tree). This means that the action potential can affect a larger area of its target cell, for example a muscle, than it could if it didn’t have ‘roots’. Just as dendrites have specific points of contact called synapses, where they receive information from the environment or other cells, so too do axons. Axons form synapses with muscles, glands, or when located deep within a network of the CNS, with other neurons (Figure 5). In fact the synapse actually contains both the transmitting point of contact (axon) and the receiving point of contact (dendrite). The cell transmitting the signal is called the presynaptic cell for before the synapse, whereas the cell receiving the signal is the postsynaptic cell for after the synapse. Our neurons are classified into two main groups depending on what other cells they make connections with and what type of information they convey. Neurons that receive input from the environment, and transmit that input into the CNS are called sensory neurons. Whereas neurons that carry information out of the CNS and make connections with muscles and glands are called motor neurons. If we are going to be able to understand how neurons make functional networks it is going to be very important to understand exactly how the neurons connect together. Presynap)c cell Axon terminal Synap)c cle4 Postsynap)c cell Figure 5: Synapse. The end of the axon divides into fine branches that swell to form axon terminals. These axon terminals are separated from the postsynaptic cell by the synaptic cleft. ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ At a synapse, the cell sending a signal is called what? (Hint: It’s the cell before the synapse.) ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ At a synapse, the cell receiving the signal is called what? (Hint: It’s the cell after the synapse.) ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ 41 LESSON READING DEFINITIONS OF TERMS Within the depths of the network in the central nervous system, neurons connect to other neurons, so the presynaptic site is usually on an axon and the postsynaptic site is usually a dendrite (we may run into exceptions later, but for now don’t worry about them). The points of contact on the axon are specialized swellings on the axon’s branches called axon terminals or presynaptic terminals, while the points of contact on the dendrite are called, not surprisingly, postsynaptic terminals. It is an important characteristic of synapses that the pre- and postsynaptic terminals do not physically touch each other. Instead, they are separated by a space called the synaptic cleft. In order to get the signal across the synaptic cleft, and depolarize or hyperpolarize the dendritic membrane the presynaptic terminal turns the action potential into a chemical signal that can cross the physical space. We will talk about this process of transmitting a signal across the synapse called synaptic transmission in another lesson. Axon terminals/Presynaptic terminals – swellings at the end of the axon’s branches that serve as the transmitting site of the presynaptic cell. As you might imagine, the function of the neuron critically depends on how long its axon is. Neurons with long axons are able to convey information over long distances to your big toe and so are called projection or relay neurons. Neurons with short axons are only able to convey information into a limited region and integrate information within a specific local area. Synaptic cleft – small gap in the synapse that separates the presynaptic cell from postsynaptic cell. So now we can classify neurons into three groups on the basis of their function: For a complete list of defined terms, see the Glossary. Wo r k b o o k Lesson 2.1 Neuronal function • Sensory neurons carry information into the central nervous system for perception. • Motor neurons carry commands out of the central nervous system to muscles and glands. • Interneurons carry information from area to area within the nervous system. They are by far the largest class, consisting of all the neurons that are not specifically sensory or motor. In summary, although all neurons contain the same three functional components, they do not all look or behave the same (Figure 6). Figure 6: Examples of neurons. Neurons that perform different functions have different shapes. Sensory neurons receive input from a sensory organ, like the ear. Motor neurons control muscle information. Local interneurons integrate activity within a small area. Projection neurons convey information for long distances. Neuroendocrine cells release hormones into blood vessels. Model neurons show that each of the different types have the same functional components. What is the name of the site on the axon that connects to the dendrite? ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ What is the name of the site on the dendrite that connects to the axon? ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ Are the axons and dendrites in physical contact with each other? ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ ___________________________________ 42 STUDENT RESPONSES How are neurons specialized to complete their functions? _____________________________________________________________________________________________________ ____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ Remember to identify your sources _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ Given what you know about the different types of neurons, what types of neurons do you predict to be involved in your ability to smell warm chocolate chip cookies? And then taste one after you eat it? What has to happen after you’ve smelled the cookie, but before you make the first bite? Be as specific as you can. ____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ _____________________________________________________________________________________________________ Wo r k b o o k Lesson 2.1 _____________________________________________________________________________________________________ ________________________________________________________________________________________________ ________________________________________________________________________________________________ 43