Signal reception
- Key People:
- Karl von Frisch
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The receiver’s task is to detect signals against the background and to discriminate between different signals. Most animals use the same sense organs (eyes, ears, noses, touch receptors, etc.) for signals that they use to detect other external stimuli. Their brains also process all sensory stimuli, both signals and nonsignals, with similar procedures. However, communication is sufficiently important that most animals show some tuning of their sense organs and some specialization of their brains to improve signal detection and characterization.
Detection of sound is often challenging because the received signals are faint and distorted owing to propagation. Sound traveling in air is largely reflected from solid objects, including animals, with little energy transfer. Sound traveling in water is easily transferred to aquatic animals, but because all parts of the animal vibrate in synchrony, there is no immobile reference allowing the animal to detect the vibrations. As a result, animals have had to acquire some very sophisticated adaptations to hear sounds.
Terrestrial animals often have funnel-shaped structures outside the body to collect and concentrate impinging sounds. The funnel shape also creates a gradual change in the properties of the sound-propagating medium from that of air to that of liquid and solid bodies. This increases the amount of trapped sound energy. At the end of the funnel is a thin membrane (called an eardrum) that is set into vibration by the sounds. Small bones or fibres transfer the eardrum movements to a fluid-filled cavity, within which are sensory cells bearing hairlike cilia. As the fluids move, the cilia sway, thereby stimulating the attached nerve fibres. Ears of orthopteran insects (e.g., crickets and grasshoppers) and vertebrates are designed such that different frequencies of sound stimulate different sets of sensory cells. Thus, the animal is able to decompose a complex sound into its component frequencies. This enables the receiver to separate signal frequencies from those of nonsignal sounds. For each band of useful frequencies, amplitudes and a temporal pattern are encoded in the rates and patterns of nerve impulses sent to the brain. Larger animals usually have two ears and use the time delays or differences in signal amplitude at the two ears to identify the direction of the sender. Small animals cannot use this method because the delays and intensity differences are very small. However, they can achieve some directional information by sampling a sound field at several points simultaneously. For example, crickets have their eardrums on their legs, and fine tubes connect each eardrum internally to breathing holes on the sides of their bodies. This allows them to obtain multiple samples of a given sound field at each point in time. Frogs and birds have a space connecting their eardrums that works in a similar manner.
Some aquatic animals have exterior cilia or hairs that sway as sounds pass over them and stimulate sensory cells. This mechanism is effective only within a few wavelengths of the sound source and tends to be limited to lower frequencies. For sound reception at a greater distance from the source, an aquatic animal must create body parts that are moved in different ways during the passage of sound waves. Some bony fish use swim bladders for this purpose. These are air-filled sacs that provide buoyancy. Because air is much more compressible than water, sound trapped by the swim bladder results in much larger molecular displacements than the same sound energy propagating in the rest of the fish’s body. Connecting the swim bladder to the fish’s ears makes it possible for the differences in molecular displacement between the swim bladder and the rest of the fish to be used to stimulate the sensory cells of the ear. This allows a fish far from the sender to detect and measure the passing sound waves.
Animal eyes differ markedly in their range of view, their resolution, and their focusing power. The eyes of vertebrates and cephalopods (octopus and squid) have adjustable lenses that extend the range over which images are in focus. They also have an iris that adjusts the amount of light entering the eye. An effective eye must have many receptor cells if it is to preserve the relative positions of the different objects reflecting light to it. To discriminate between different colours, different receptor cells must be sensitive to different light frequencies. In addition, because light intensities are highly variable, eyes may need different receptor cell types to handle both dim and bright light. Vertebrate eyes use cone receptor cells for bright light and colour discrimination and colour-insensitive rod receptors for dim light conditions. All light receptors contain a protein-bound pigment. This pigment, called rhodopsin, occurs in all multicellular animals. These pigment molecules change shape when absorbing light. This triggers a chain of reactions within the receptor cells ending in the production of nerve impulses. Different photopsins and rhodopsins absorb different light frequencies, permitting receptors to differ in colour sensitivity. Arthropod eyes consist of 8–10 receptor cells clustered around each of many facets, or corneal lenses. It is the number of facets, not the number of receptor cells, that determines visual acuity in arthropod eyes.
Unlike the case with auditory signals, the direction of a visual sender is automatically known once a receiver has detected the signal. The relative positions of receptor cells in an eye are preserved in their projections to the brain. This allows the brain to create a map that replicates, to varying degrees, the visual field of the eye. Projections from the visual maps of the two eyes are compared to identify specific objects and their relative distances. The recognition of patterns is a function of the brain and relies on a combination of inherited and learned mechanisms.
Pheromone reception is accomplished by smell (olfactory) or taste (gustatory) organs. These receptors contain sensory cells with fine cilia, or microvilli, that extend into the medium. Pheromone molecules and other stimuli temporarily bind to specific proteins on the cilia or microvilli. The binding triggers a cascade of chemical reactions within the sensory cell that result in the production of nerve impulses flowing to the brain. The more molecules that bind to a cell, the higher the rate of nerve firing. The sensory cells in an olfactory organ can have highly specific or generalized sensitivities. Specific cells tend to respond only to a certain molecule. Animals often use such cells to detect species-specific mating pheromones. Generalized cells respond to several types of molecules. They are used for less-specific functions, such as the recognition of food items. The olfactory organs of different species vary in the mix of specialized versus generalized sensory cells that they host. Olfactory organs can be as simple as a patch in a mammalian breathing passage or as complex as the plumose antennae of male moths. In some mammals the vomeronasal organ (Jacobson’s organ), located in the roof of the mouth, is used to mediate a behavioral response known as flehmen, in which an animal raises its head and lifts its upper lip in reaction to specific odours. This response requires special movements of the tongue and lips to admit chemical samples to the sensory cells. The vomeronasal organ is the primary receptor organ for many of the pheromones that dictate interactions in mammalian social life, including pheromones involved in conflict, reproduction, and parental care.
Electroreception appears to have been a widespread sensory ability in primitive fishes, in which it was used to detect muscle and nerve impulses in hidden prey. Modern sharks and rays still use this technique for hunting. Mormyriform and gymnotiform electric fish developed these primitive receptors into sophisticated tuberous organs that are used in social communication. These structures, embedded in the fishes’ skin, are encapsulated in ways that make them insensitive to slowly varying electric fields, such as those produced by muscles, but responsive to the rapid discharges of other electric fish. Tuberous receptors are usually tuned to be most responsive to the discharge rate of their own species. The brains of these fish are also highly sensitive to changes in the repetition rates of discharges. This permits sophisticated exchanges between fish during conflicts, courtship, and territorial defense.
Costs and benefits of communication
For both senders and receivers there are costs associated with engaging in communication. It takes time, energy, and special modifications of sender and receiver organs to communicate. Thus, there must be compensatory benefits to each party for communication to be favoured by evolution. A sender will provide information to a receiver only if the decision of the receiver improves the sender’s fitness more than the costs of signaling reduces it. The benefits to the sender may be direct, such as securing a mate or successfully repelling an opponent, or indirect, in that the receiver’s choice may benefit close kin of the sender. A receiver attends to any source of information that is sufficiently reliable, on average, to enhance the receiver’s decision making. The qualifiers “sufficiently reliable” and “on average” reflect the fact that senders may not always send perfect information, but the signals may still be useful to receivers. However, there is a minimum amount of reliable information in any decision situation that must be provided before it is beneficial for either party to engage in communication. Described another way, the payoff to each party (benefits minus costs) must be positive before they will participate in communication.
Animals often have to make decisions in response to alternatives. For example, if a female must determine which of two males will be the better parent for her young, she will mate with the male she deems most fit. In another example, when a parent bird returns to its nest, it must decide which of its nestlings is most needy. It will then give that nestling the worm. The receiver’s decision process begins with some baseline probabilities for each alternative, based on inherited biases, chance, or prior experience. As new information is obtained, whether by examining the candidates directly or by attending to their signals, the receiver updates the probabilities by raising some and lowering others. The receiver’s decision will then be based on these updated probabilities and some knowledge of the relative payoffs of correct versus incorrect decisions.
A number of measures have been proposed for the amount of information provided by animal signals. One measure is relative reliability; this is the increase in the average probability that a receiver makes the correct decision (for itself) when using signals compared to when a receiver is not using signals. Relative reliability also may be weighted according to the relative chances of making the correct decision; this is done by determining the difference in payoffs when making the correct versus incorrect decision. A classical approach computes a logarithmic measure (in bits) of the reduction in a receiver’s uncertainty after receiving and processing a signal. Most of these measures yield similar rankings of the amount of information in signals and differ largely in scale.
Few senders never err in signal selection, and even if they selected the correct signal, transmission can distort signals and cause receivers to interpret them incorrectly. Thus, most animal signaling systems have reliabilities less than that expected for perfect information. Receivers usually have a “best guess” that they adopt when they cannot obtain enough information from a signal to make an informed choice. This best guess is the one that yields the highest average payoff when no signals are available. A receiver that switches from best guesses to a reliance on imperfect signals will start making incorrect decisions, when the best guess was in fact the correct choice. However, the receiver will also start making some correct decisions on those occasions when the best guess was the incorrect choice.
For receivers to participate in communication, the benefits of not making errors when the best guess was wrong must be greater than the costs of occasional errors when the best guess was always correct. As reliability increases, these benefits keep growing and the costs become diminishingly small. Thus, there is a minimal reliability that is required before either party should engage in communication.
