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Major Question in Marine Biology: How do social systems help regulate sex change in fishes? The behavior of sex change is affected by the social environment, and requires the reorganization of physiological systems. From gonads being transformed, to changes in body size, these physical changes happen as a fish take cues about the size of conspecifics and their behavior from surrounding social environment. These social behaviors help regulate reproduction and growth rate. The composition of a social group triggers the sex change through encounters with smaller and larger fish, and inhibits change by the continued presence of a large fish conspecific. Hermaphroditic fish function as both male and female either sequentially or simultaneously for some period during their life. The following three papers answer the question of how social systems help regulate sex change in fish through experimentation and by establishing criteria for diagnosis of hermaphroditism in fish. They are summarized below. There are a wide variety of criteria and methods for assessing whether a fish species is a hermaphrodite. After the discovery that sex change in sequential hermaphrodites can be induced by social cues, many various scientists began work on the behavioral, ecological, and evolutionary aspects of hermaphroditism. Many methods and criteria used to diagnose hermaphroditism in older studies needed to be reassessed in light of the knowledge we have today. Thus, the authors set out to determine whether a fish is hermaphroditic. (Sadovy and Shapiro, 1987) Some of the features typically used to establish hermaphroditism are more reliable than others. In addition, there are three forms of hermaphroditism. First, conversions from female to male are known as protogynous. Second, conversions made from male to female are called protandrous, and are less common. The third form, called simultaneous hermaphroditism, occurs when the fish has both male and female organs. Protogyny is strongly suggested by the physical make-up of a reproductive system, the presence of transitional individuals, and through manipulating the social system to experimentally produce sex reversed fish. Bimodal size and age frequency distributions are less reliable as this could result from many causes, only one of which is protogyny. Protandry is more challenging to diagnose because testicular tissues from the previous state of sex are usually gone. Detecting protandry is best done by performing an experimental behavioral induction of sex change as age and size frequency distributors are unreliable indicators of this condition. Simultaneous hermaphroditism is diagnosed by the presence of mature tissue from both sexes in one gonad followed by inducing self-fertilization. Large sample sizes are required to diagnose hermaphroditism in fish, including individuals of all sizes, throughout the year from a single location. The only known way to cause complete sex change is by removing a female, which causes a male to change sex, or by removing a male which causes a female to change sex. This must also be coupled with histological examination. There are specific guidelines that should be followed when performing a sex change induction experiment. Social conditions should resemble nature as closely as
possible, and there should be a large sample size. Last, but not least, the sex of each fish should be determined before and after the experiment. (Sadovy and Shapiro, 1987) How is sex change socially controlled in non-harem fish species with less rigid social and mating systems? Fish species that are harem-living or have rigid dominance hierarchies exhibit social control of sex change which happens rapidly with social status, and is reversible. Usually, this is due to loss of a dominant male or female, which causes sex change in the dominant fish of the opposite sex. In non-harem species, sex change is more flexible. The saddleback wrasse, Thalassoma duperrey, a non-harem species, was used to show sex change in these fish is socially controlled, and is instigated by visual stimuli. This coral reef species exhibits protogynous hermaphroditism, and lives in sexually integrated home ranges. Rather than mating in a harem, promiscuous mating occurs. Sex change in this species is a function of relative sizes in the social group. The initiation of sex change in individual females is due to the proportion of larger or smaller fish within the home range. The largest female living in a mate’s territory will make the sex change within 2-3 weeks after the mate disappears. The larger female adopts the male sexual behavior to mate with smaller females. The bigger a fish, the better it can defend the territory. Therefore, the largest female assumes the role as male. Interestingly, behavioral sex change is reversible in these fish as when a male is reintroduced after a female sex change, the sex-changed female will accept the male as her mate. (Ross and Losey, 1983) These fish change quickly with changes in social status in their social system. In other words, there are fish that don’t stop with one sex change. Instead, the amount of gender changes depends on the stability of the environment of the social system. Behavior of the fish is a good indicator of male or female status. For example, males not only behave aggressively by defending their nests, but they also perform a dance for their chosen mate. Relative size was crucial whereas coloration was not. Optimal reproductive success depends on the relative numbers of larger and smaller fish. If the proportion of large fish is low, sex change would be a good solution. Consequently, the mechanism of size-ratios used by T. duperrey appears to be a good solution as they live in a socially variable environment. These males show social control of sex change because dominant males inhibit sex change in other fish in the social system by being aggressive. These fish regulate the amount of reproduction within their group. Social control over sex change occurs, and allows a response to social cues. This is an advantage in heterogeneous environments like coral reefs. (Ross and Losey, 1983) What are the early behavioral and morphological changes associated with dominance and sex change? Early behavioral and morphological changes associated with dominance and sex change are socially controlled. Inhibition by males and stimulation from other females are the two social features that regulate sex change. In May and June of 2002, the fish species, Ltthrypnus dalli, known as blue-banded gobies, were collected around Santa Catalina Island off the coast of California, and two experiments were performed. (Rogers, Drane, et. al. 2005) Interdependence of behavior between males and females was displayed as the behavior of females had a direct impact on male behavior rates. The first experiment quantified the morphological traits and behavioral interactions between females. Experiment two demonstrated that dominant fish show disproportional growth in their dorsal fin. With blue-banded gobies, a pair of females is a large enough group to create a
sex change. The dominant fish showed male courtship behavior, was the instigator of most of their encounters, and initiated spawning behavior. When pairing two female blue-banded gobies, the dominant fish became male. One fish became dominant over the other, and began showing behavioral and morphological characteristics of males. The other fish remained female, showing sexual behaviors of females and morphological characteristics of females. In, blue-banded gobies, sexual dimorphism occurs as the males are longer. When one female becomes dominant, elongation of the dorsal fin is the first morphological change noticed followed by rapid overall growth of the fish. All of the dominant fish showed male gonad morphology and the subordinate fish clearly had eggs present within the ovaries. In addition, the subordinate fish behaved more passively, like females as when a dominant fish approached, the subordinate fish was displaced. In conclusion, these fish base their behavior on the behaviors of other fish in their environment. (Rogers, Drane, et. al. 2005) The most recent paper relates to the original paper about criteria for sex change in fish because the experiment follows the correct procedures. This experiment, like the experiment in the earlier paper, looks at protogynous fish. Again, confirming that sex change is socially controlled. The sex change is a direct response to changes in social structure. The two social features that regulate sex change are inhibition by males, and stimulation from other female fish. Early behavioral and morphological changes are associated with dominance and sex change. For example, in a pair of females, the dominant fish changed sex from female to male. This study shows that males are larger, showing sexual dimorphism. The original paper addresses sexual dimorphism or the fact that sex change in fish affects the behavior and morphological characteristics of males. However, an interdependent relationship between male fish and female fish was shown in this study. The behavior of sex change is affected by the social environment which causes fish to reorganize their physiological systems. These social behaviors of fish regulate successful reproduction and their growth rate. CONCLUSION: Unanswered questions Although several questions were answered in the discussion, there are still questions about sex change in fish that remain unanswered. How do fish know that another fish is dominant the day after a confrontation? There is most likely not memory involved, but just the aggressive day to day behavior of the dominant male at work. Why can some fish change sex in fewer than 3 days, while others take up to several years? This question is possibly answered by the fact that nature has a way of acting when necessary. For example, in the case of simultaneous hermaphrodites, who swim in the deep sea. The low light levels along with a shortage in food limits the fish population. Therefore, mating partners are hard to find so fish living there have to be able to mate with whatever fish of their species comes along, male or female. Amazingly, nature has provided fish with this ability in order to keep reproducing in their challenging environments.
Bibliography Ross RM, Losey GS, Diamond M. 1983 Jan 31. Sex Change in a Coral-Reef Fish: Dependence of Stimulation and Inhibition on Relative Size. Science [serial on the internet]. 221 No. 4610: 574 – 575. Available from: http://www.sciencemag.org/cgi/content/abstract/221/4610/574 Sadovy Y, Shapiro DY. 1987 Feb. Criteria for the diagnosis of hermaphroditism in fishes. Copeia [serial on the Internet]. No. 1: 136-156. Available from: http://www.jstor.org/browse/00458511/ap050402/05a00170/0? frame=noframe&userIDfirstname.lastname@example.org/01cc99331200501b7cb36&b ackcontext=page&backurl=/cgibin/jstor/viewitem/00458511/ap050402/05a001 70/0%3fframe%3dnoframe%26dpi%3d3%26userID %email@example.com/01cc99331200501b7cb36%26config%3djstor %26PAGE%3d0&config=jstor Rodgers EW, Drane S, Grober MS. 2005 April. Sex reversal in pairs of Lythrypnus dalli: behavioral and morphological changes. Biol. Bull. [serial on the Internet]. 208: 120-126. Available from: http://www.biolbull.org/cgi/content/abstract/208/2/120
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