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The Evolution of Multiple Sensory Modalities Used on Kin Recognition
Department of Biology
Lake Forest College
Lake Forest, Illinois 60045
Kin recognition is an important regulator of animal behavior in nature. It is used for several reasons, including aggression, mating, and parental care. Various animals use multiple sensory modalities for social interactions in the environment. Kin recognition has multiple impacts on species due to the use of several sensory modalities which include physiological changes, imprinting and neural development. This paper will focus on three organisms which use several sensory modalities in kin recognition: the zebrafish, the clonal fish, and the common house mouse. Each of these organisms uses several sensory modalities reasons which including imprinting, parental care, and regulation of aggression. Each example reveals the importance of the social interactions caused by the multiple sensory modalities used during kin recognition. Additionally, they expand on how this has impacted the evolution of the nervous system of animals and the adaptions in animal behavior.
One of the most important aspects of social behavior in nature is kin recognition. This is a process in which a related group within a species can identify who they are related to in nature. Kin recognition has influenced how animals communicate and act within their environment (Hinz et al., 2013). It is important for several reasons, which include prevention of inbreeding, regulation of aggression, and improving energy investment of the parents (Longru et al., 1983, Makowicz, 2016).
Kin recognition involves several behavioral and physiological changes of the parent and the offspring during development. It may also involve changes in specific genes that influence kin recognition (Aken, 2009). For example, maternal and paternal mice have a gene called olfactory receptor 692 (Olfr692), and the neurons expressed by this gene are activated when the pup odors encounter the parent’s olfactory system to create instinctive actions towards young (Nakahara, 2016).
Kin recognition uses several mechanisms to create recognition within species. Several of these mechanisms include location, territory, social learning, and phenotypic matching (Blaustien, 1983). These mechanisms of kin recognition require the use of the various sensory modalities of vision, olfactory, and auditory senses (Hinz, 2013). Many species use olfactory senses in addition to one of the other sensory modalities in kin recognition (Coffin, 2011). Olfactory senses are commonly used in kin recognition because they often reveal the “genetic link” to other kin (Dubas, 2009). The use of several sensory modalities is very important for mother-child bonding and kin recognition for imprinting (Hinz et al., 2013).
Imprinting during infancy results in kin recognition and sometimes requires several sensory modalities to ensure that no mistakes are made in recognition of stimuli within an environment (Hinz et al., 2013). Imprinting allows the mother to distinguish her young from others and it allows the young to recognize their mother (Longru et al., 2015). In many species, mothers use several sensory modalities to identify and locate infants through “multisensory integration of the nervous system”. This is important to get attention and elicit a response from the mother (Cohen 2011). The function of kin recognition regarding several sensory modalities is a relatively new study that is slowly gaining popularity. It is important to understand how multiuse of sensory systems within an environment explains the physiological and genetic evolution within species that influence social behavior.
Visual and Olfactory Sensory Modalities of Zebrafish
In this experiment, researchers were trying to determine if kin recognition in zebrafish, Danio rerio, requires both olfactory and visual sensory integration for successful imprinting (Hinz et al., 2013). The methods of this experiment included raising zebrafish larvae and exposing the larvae to different visual and olfactory cues 12 days after fertilization (Hinz et al., 2013). Olfactory cues were given 5-7 days after fertilization by placing groups in kin water or non-kin water (Hinz et al., 2013). To see if visual stimuli was required for imprinting, the experimenters showed images of kin to various groups of zebrafish on day 6 with various conditions, with and without olfactory cues (Hinz et al., 2013). At days 8-12 post fertilization, the odor choice test was used in an Atema flume to see whether the different experimental groups of guppies preferred kin or non-kin water (Hinz et al., 2013). To measure the preference of the larvae the experimenters recorded which side of the flume they swam to (Hinz et al., 2013).
The results provided show that the larvae preferred kin water (Hinz et al., 2013). However, for the larvae to recognize kin, both olfactory and visual were needed, which is shown in the group OkVnk (Hinz et al., 2013). Having olfactory or visual cues alone had low success rates in imprinting on the zebrafish larvae (Hinz et al., 2013).
Auditory and Olfactory Sensory Modalities of Mice
In this experiment, the experimenters were trying to understand how imprinting influences the actions of female mice, Mus musculus, and if this requires the use of several sensory modalities (Cohen, 2011). The two female groups used consisted of maternal and virgin mice which were exposed to mice pups (Cohen, 2011). The experimenters recorded neuron activity in the A1 region (auditory cortex) of the brain of maternal and virgin female mice when pup noises were presented to them(Cohen, 2011). While the pup noises were being used, odors coming from the pups were filtered to the monitored female mouse (Cohen, 2011).
The results show that the maternal mice had the highest stimulation of neuronal activity in A1 with pup noises and odor compared to virgin females (Cohen, 2011). A lower response rate to pup noises was recorded in mothers without olfactory stimulation (Cohen, 2011). It can be deduced that the neurons in the A1 region of the brain required the integration of both olfactory and auditory sensory cues in maternal mice to create instinctive reactions (Cohen, 2011).
Olfactory and Visual Sensory Modalities of Clonal Fish
Poecilla formosa, or amazon mollies, reproduce asexually. As a result of this, they are considered clonal fish; however, they do have small genetic variation between generations. Offspring can be up to 50% genetically identical to other siblings (Makowicz, 2016). The experimenters in this study were trying to understand how Clonal fish distinguish kin from non-kin with small genetic variance and whether they use several sensory modalities (Makowicz, 2016). Clonal and nonclonal sister groups were used in the experiment (Makowicz, 2016). The different aggressive interactions between siblings and non-kin were observed through tanks that had different sensory stimuli exhibited through glass and tubes (Makowicz, 2016). In different treatments, chemical, visual, and mechanical cues were passed between the groups through tubes (Makowicz, 2016). Additionally, an open water environment was stimulated for the sister and non-sister groups and aggressive behaviors were observed (Makowicz, 2016).
The figure provided shows that visual, chemical, and mechanical sensory modalities are used in kin recognition for clonal fish (Makowicz, 2016). Clonal fish that were related to one another spent more time with each other and had less aggressive behavior than with fish they were not related to (Makowicz, 2016). Additionally, in this case multiple sensory modalities weren’t needed for kin recognition (Makowicz, 2016). However, these sensory modalities could all be used for kin recognition in clonal fish (Makowicz, 2016). A slightly higher preference was shown for visual cues compared to olfactory cues (Makowicz, 2016).
From the results of these three studies, it can be concluded that several sensory modalities are used in kin recognition for various reasons. Additionally, multisensory kin recognition is an important aspect in regulating behavior between kin and non-kin in nature. The results from the zebrafish study show that multisensory integration of visual and olfactory senses for imprinting occurs early on in development and is essential to later kin recognition The results also show olfactory and visual stimulation alone did not create kin recognition This may be because more information is needed for imprinting to help the species distinguish conspicuous stimuli from other common stimuli in the environment Olfactory cues limit the ability of the fish in the to distinguish sensory stimuli in the environment, while visual stimuli narrows the information needed by the zebrafish in the kin recognition process Zebrafish did not imprint on non-kin, further supporting the idea that kin recognition requires a level of “genetic predisposition” in the cognitive system (Hinz et al., 2013). The incorporation of multisensory integration can be seen to have evolved in species to create further distinction of kin to facilitate social interactions. (Hinz et al., 2013). This study shows that multiple sensory modalities are important for kin recognition and have evolved to create multisensory integration that influences the cognition of the animal (Cohen, 2011). Another aspect this study touches on is how time of development is important with the exposure to sensory modalities in the cognitive process of kin recognition .Other studies have found that timing of kin recognition is important because the various mechanisms involved in the neurological process of imprinting allow animals to switch from predatory to parentlike behaviors in many species (Odekop, 2007).Possible future studies could investigate how the brain interprets and processes kin recognition with sensory modalities at the beginning stages of life.
Multisensory integration is also important for parents raising offspring, due to the energy investment required for raising young (Cohen, 2011). Having additional sensory steps in kin recognition help the parents to analyze the costs and benefits of the energy investment (Hinz et al., 2013). As seen with the mice experiment, multiple sensory modalities, as well as imprinting, are needed to influence maternal responses (Cohen, 2011). It can be concluded that auditory cues are used to find the mice pup and olfactory cues are used for pup identification (Cohen, 2011). The results also show that brain activity in maternal and virgin mice differ and that multisensory integration is required to fully elicit a response from the mother (Cohen, 2011). The difference in response between mothers and virgins in the A1 region of the brain shows that, “plasticity is experience dependent” (Cohen, 2011). The plasticity of senses from the mother are a result of physiological changes, exposure to infant smell, and going through the experience of motherhood (Cohen, 2011). Parental care using several sensory modalities regarding kin recognition is important for reproductive success and energy investment (Mathews, 2010). Maternal instincts are especially important because this energy investment is put into the protection of infants, which increases the reproductive fitness of the mother (Mathews, 2010). The multisensory integration of parents prevents them from caring for the wrong offspring and gives them an evolutionary advantage in reproductive fitness (Nakahara, 2016). An interesting study to further this research would be to see whether the victims of species due to brood parasites are due to the evolutionary lag in multisensory integration.
Clonal fish expressed the ability to distinguish kin from non-kin through sensory modalities, regardless of ha having only minimal differences between offspring (Makowicz, 2016). The results support the statement that kin recognition has evolved and adapted to genetic similarities by using various sensory modalities to distinguish kin (Makowicz, 2016). The results show that integration of different sensory modalities is not always used (Makowicz, 2016). However, although multiple sensory modalities are not always required for kin recognition, they may be important for sensory plasticity in different environments (Stevens, 2013). The experimenters only looked at how the fish interacted during the day and not at night in clear water conditions (Makowicz, 2016). At later times of the day, certain species could end up using multisensory kin recognition to identify kin.
Even in a genetically similar environment, lower aggression was seen towards kin .This shows that multisensory kin recognition is an important evolutionary aspect of animal behavior (Makowicz, 2016). Being able to recognize kin allows parents to better protect their offspring against predators and to allocate them food resources; this behavior can be seen in many species, including crayfish (Mathews, 2010). Kin recognition has also been shown to influence where penguins nest and seek shelter (Coffin, 2011). This all shows that kin recognition is a significant factor in the evolution of animal behavior and interactions within a species.
Summary and Conclusion
As demonstrated, kin recognition is very important in social interactions within the environment and has evolved to use several sensory modalities. Several sensory modalities are used in kin recognition to regulate parental care, aggression, mating, and imprinting. To reduce incorrect behavior responses to kin based sensory stimuli, multiple sensory modalities are imperative to evolutionary development within species. Sensory modalities have evolved and become integrated within the nervous system of certain species and more research is needed to understand how integrated sensory modalities influence kin recognition and animal physiology in nature.
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