What Is the Difference Between a Graded Potential and Action Potential and How Does This Impact Impulse Transmission?
There are important differences between graded potentials and action potentials of neurons (see Introduction to this lecture). Table 1 lists the main differences between graded potentials and action potentials. As discussed in this lecture and upcoming lectures, most of these differences are due to the fact that graded potentials result from the passive electrical property of the neuronal membrane, whereas action potentials result from an orchestrated response to depolarizing stimuli, and involve a coordinated activity of voltage-gated ion channels. Graded potentials must occur to depolarize the neuron to threshold before action potentials can occur.
Important insights into the nature of electrical signals used by nerve cells were obtained 80 years ago. Electrodes were placed on the surface of an optic nerve of an invertebrate eye. (By placing electrodes on the surface of a nerve, it is possible to obtain an indication of the changes in membrane potential that are occurring between the outside and inside of the nerve cell.) Then 1-sec duration flashes of light of varied intensities were presented to the eye; first dim light, then brighter lights. Very dim lights produced no changes in the activity, but brighter lights produced small repetitive spike-like events. These spike-like events are called action potentials, nerve impulses, or sometimes simply spikes. Action potentials are the basic events the nerve cells use to transmit information from one place to another. The recordings in the figure above illustrate three very important features of nerve action potentials. First, the nerve action potential has a short duration (about 1 msec). Second, nerve action potentials are elicited in an all-or-nothing fashion. Third, nerve cells code the intensity of information by the frequency of action potentials. When the intensity of the stimulus is increased, the size of the action potential does not become larger. Rather, the frequency or the number of action potentials increases. In general, the greater the intensity of a stimulus, (whether it be a light stimulus to a photoreceptor, a mechanical stimulus to the skin, or a stretch to a muscle receptor) the greater the number of action potentials elicited. Similarly, for the motor system, the greater the number of action potentials in a motor neuron, the greater the intensity of the contraction of a muscle that is innervated by that motor neuron. Action potentials are of great importance to the functioning of the brain since they propagate information in the nervous system to the central nervous system and propagate commands initiated in the central nervous system to the periphery. Consequently, it is necessary to understand thoroughly their properties. To answer the questions of how action potentials are initiated and propagated, we need to record the potential between the inside and outside of nerve cells using intracellular recording techniques.The potential difference across a nerve cell membrane can be measured with a microelectrode whose tip is so small (about a micron) that it can penetrate the cell without producing any damage. When the electrode is in the bath (the extracellular medium) there is no potential recorded because the bath is isopotential. If the microelectrode is carefully inserted into the cell, there is a sharp change in potential. The reading of the voltmeter instantaneously changes from 0 mV, to reading a potential difference of -60 mV inside the cell with respect to the outside. The potential that is recorded when a living cell is impaled with a microelectrode is called the resting potential, and varies from cell to cell. Here it is shown to be -60 mV, but can range between -80 mV and -40 mV, depending on the particular type of nerve cell. In the absence of any stimulation, the resting potential is generally constant.
Information is encoded in neural circuits using both graded and action potentials, converting between them within single neurons and successive processing layers. This conversion is accompanied by information loss and a drop in energy efficiency. We investigate the biophysical causes of this loss of information and efficiency by comparing spiking neuron models, containing stochastic voltage-gated Na+ and K+ channels, with generator potential and graded potential models lacking voltage-gated Na+ channels. We identify three causes of information loss in the generator potential that are the by-product of action potential generation: (1) the voltage-gated Na+ channels necessary for action potential generation increase intrinsic noise and (2) introduce non-linearities, and (3) the finite duration of the action potential creates a ‘footprint’ in the generator potential that obscures incoming signals. These three processes reduce information rates by ∼50% in generator potentials, to ∼3 times that of spike trains (Koch C, Douglas R, Wehmeier U, 1990). Both generator potentials and graded potentials consume almost an order of magnitude less energy per second than spike trains. Because of the lower information rates of generator potentials they are substantially less energy efficient than graded potentials. However, both are an order of magnitude more efficient than spike trains due to the higher energy costs and low information content of spikes, emphasizing that there is a two-fold cost of converting analogue to digital; information loss and cost inflation. As in electronics, many of the brain's neural circuits convert continuous time signals into a discrete-time binary code. Although some neurons use only graded voltage signals, most convert these signals into discrete-time action potentials. Yet the costs and benefits associated with such a switch in signalling mechanism are largely unexplored. We investigate why the conversion of graded potentials to action potentials is accompanied by substantial information loss and how this changes energy efficiency. Action potentials are generated by a large cohort of noisy Na+ channels. We show that this channel noise and the added non-linearity of Na+ channels destroy input information provided by graded generator potentials. Furthermore, action potentials themselves cause information loss due to their finite widths because the neuron is oblivious to the input that is arriving during an action potential. Consequently, neurons with high firing rates lose a large amount of the information in their inputs. The additional cost incurred by voltage-gated Na+ channels also means that action potentials can encode less information per unit energy, proving metabolically inefficient, and suggesting penalisation of high firing rates in the nervous system (Manwani A, Koch C, 1999).
As has been noted, most animal cells can fire or create their own action potential. Muscle cells contract when they fire and are often induced to do so by a nerve impulse. In fact, nerve and muscle cells are physiologically similar, and there are even hybrid cells, such as in the heart, that have characteristics of both nerves and muscles. Some animals, like the infamous electric eel use muscles ganged so that their voltages add in order to create a shock great enough to stun prey. Propagation of a nerve impulse down a myelinated axon, from left to right. The signal travels very fast and without energy input in the myelinated regions, but it loses voltage. It is regenerated in the gaps. The signal moves faster than in unmyelinated axons and is insulated from signals in other nerves, limiting crosstalk.
Manwani A, Koch C (1999) Detecting and estimating signals in noisy cable structure, I: neuronal noise sources. Neural Comput 11: 1797–1829
Aguera y Arcas B, Fairhall AL, Bialek W (2003) Computation in a single neuron: Hodgkin and Huxley revisited. Neural Comput 15: 1715–1749
Lenn NJ, Reese TS (1966) The fine structure of nerve endings in the nucleus of the trapezoid body and the ventral cochlear nucleus. Am J Anat 118
Rekling JC, Funk GD, Bayliss DA, Dong XW, Feldman JL (2000) Synaptic control of motoneuronal excitability. Physiol Rev 80: 767–852
Koch C, Douglas R, Wehmeier U (1990) Visibility of synaptically induced conductance changes: theory and simulations of anatomically characterized cortical pyramidal cells. J Neurosci 10