Sequential hermaphroditism (called 
dichogamy in 
botany) is a type of 
hermaphroditism that occurs in many 
fish, 
gastropods  and plants. Here, the individual is born one sex and changes sex at  some point in their life. They can change from a male to female (
protandry), or from female to male (
protogyny).
[1]  Despite which sex the organism changes to, those that change gonodal  sex can have both female and male germ cells in the gonads or can change  from one complete gonodal type to the other during their last life  stage.
[2]
Zoology
Protandry
Protandry refers to organisms that are born male and at some point in  their lifespan change sex to female. Protandrous animals include 
clownfish. Clownfish have a very structured society. In the 
Amphiprion percula species, there are zero to four individuals excluded from breeding and a breeding pair living in a 
sea anemone.  Dominance is based on size, the female being the largest and the male  being the second largest. The rest of the group is made up of  progressively smaller non-breeders, which have no functioning gonads.
[3]  If the female dies, the male gains weight and becomes the female for  that group. The largest non-breeding fish then sexually matures and  becomes the male of the group.
[4]
Other examples of protandrous animals include:
- The ctenophore Coeloplana gonoctena.  In this organism the females are bigger than the males and are only  found during the summer. In contrast males are found year round.
- The flatworms Hymanella retenuova and Paravortex cardii.
- Laevapex fuscus, a gastropod,  is described as being functionally protandric. The sperm matures in  late winter and early spring, and the eggs mature in early summer, and copulation  occurs only in June. This shows that males cannot reproduce until the  females appear, thus why they are considered to be functionally  protandric.[5]
[edit] Protogyny
  
  Moon wrasse, 
Thalassoma lunare, a protogynous animal species
  Protogyny refers to organisms that are born female and at some  point in their lifespan change sex to males. Common model organisms for  this type of sequential hermaphroditism are wrasses. They are one  of the largest families of coral reef fish and belong to the Labridae  family. Wrasses are found around the world in all marine habitats and  tend to bury themselves in sand at night or when they feel threatened.[6]  In wrasses, the larger of the two fish is the male, while the smaller  is the female. In most cases, females and immature have a uniform color  while the male has the terminal bicolored phase.
[7]  Large males hold territories and try to pair spawn while small to  mid-size initial-phase males live with females and group spawn.
[8] In other words, both the initial and terminal phase males can breed; they differ however in the way they do it.
In the California Sheephead (
Pimelometopon pulchrum), a type of 
wrasse,  when the female changes to male, the ovaries degenerate and  spermatogenic crypts appear in the gonads. The general structure of the  gonads remains ovarian after the transformation and the sperm is  transported through a series of ducts on the periphery of the gonad and  oviduct. Here sex change is age dependant. For example, the California  sheephead stays a female for four years before changing sex.
[7]
Other examples of protogynous organisms include:
[edit] Ultimate causes
Ghiselin proposed three models for hermaphroditism in 1969 in his paper titled “The evolution of hermaphroditism  among animals”. The ‘’low-density model’’ states that individuals have  characteristics that reduce the opportunity for mating; this model  cannot be applied to sequential hermaphroditism. The ‘’gene  dispersal model’’ is based on the idea that limitations on dispersal  may influence population structure or genetical environment and it can  be separated into two versions: the inbreeding version and the  sampling-error version. This theory of gene dispersal can be applied to sequential hermaphrodites,  especially the inbreeding version. The inbreeding version is based upon  the fact that both protandry and protogyny help prevent inbreeding in  plants and thus one can make the same assumption that in animals it  works by reducing the probability of this occurring among siblings. The  sampling-error version is based on the reality that the genetical  environment is influenced by genetic drift and similar phenomena in  small populations. The two aspects of these hypotheses influenced by hermaphroditism,  that is inbreeding and sampling-error, result in the same thing,  reduction of genetic variability. In other words being a hermaphrodite  would increase genetic variability and thus be considered advantageous  to the organism. This theory of gene dispersal can be applied to sequential  hermaphrodites, especially the inbreeding version. Lastly, the  ‘’size-advantage model’’ states that reproductive functions are carried  out better if the individual is a certain size/age. Assuming that the  reproductive functions of one sex are better performed at a certain  size, then an organism would assume the sex that its size allows to  perform the best. This would increase its reproductive potential and  fitness. For example, eggs are larger than sperm, thus if you are a big  you are able to make more eggs so being female when big is advantageous,  however the size advantage relationship is really not as simple as the  example just mentioned, but it allows for a better understanding of it.[9]
In most ectotherms body size and female fecundity are positively correlated.
[1] This supports Ghiselin’s size-advantage model, which is still widely accepted today. Kazancioglu and Alonzo (2010)  performed the first comparative analysis of sex change in Labridae.  Their analysis supports the size-advantage model by Ghiselin and suggest  that sequential hermaphroditism is correlated to the  size-advantage. They determined that dioecy was less likely to occur  when the size advantage is stronger than other advantages[10]
Warner suggests that selection for protandry may occur in populations  where female fecundity is augmented with age and individuals mate  randomly. Selection for protogyny may occur where there are traits in  the population that depress male fecundity at early ages  (territoriality, mate selection or inexperience) and when female  fecundity is decreased with age, the latter seems to be rare in the  field.
[1]  An example of territoriality favoring protogyny occurs when there is a  need to protect their habitat and being a large male is advantageous for  this purpose. In the mating aspect, a large male has a higher chance of  mating, while this has no effect on the female mating fitness.
[10] Thus, he suggests that female fecundity has more impact on sequential hermaphroditism that the age structures of the population.[1]
The size-advantage model predicts that sex change  would only be absent if the relationship between size/age with  reproductive potential is identical in both sexes. With this prediction  one would assume that hermaphroditism is very common, but this is not the case. Sequential hermaphroditism  is very rare and according to scientists this is due to some cost that  decreases fitness in sex changers as opposed to those who don’t change  sex. Kazanciglu and Alonzo confirmed this in 2009. They found that the costs of changing sex only favored dioecy when the cost was very large but that some groups favored hermaphroditism. This indicates that the cost of sex change does not explain the rarity of sequential hermaphroditism by itself.[11]
[edit] Proximate causes
Many studies have focused on the proximate causes of sequential hermaphroditism.  The role of aromatase has been widely studied in this area. Aromatase  is an enzyme that controls the androgen/estrogen ratio in animals by  catalyzing the conversion of testosterone into oestradiol, which is  irreversible. It has been discovered that the aromatase pathway mediates  sex change in both directions.[12]  Many studies also involve understanding the effect of aromatase  inhibitors on sex change. One such study was performed by Kobayashi et  al. In their study they tested the role of estrogens in male three-spot  wrasses (
Halichoeres trimaculatus). They discovered that fish  treated with aromatase inhibitors showed decreased gonodal weight,  plasma estrogen level and spermatogonial proliferation in the testis as  well as increased androgen levels. Their results suggest that estrogens  are important in the regulation of spermatogenesis in this protogynous  hermaphrodite.
[13]
[edit] Botany
[edit] Flowering plants
In the context of the 
plant sexuality of flowering plants (angiosperms), there are two forms of dichogamy: 
protogyny—female function precedes male function—and 
protandry—male function precedes female function.
Historically, dichogamy has been regarded as a mechanism for reducing 
inbreeding (e.g., Darwin, 1862). However, a survey of the angiosperms found that 
self-incompatible  (SI) plants, which are incapable of inbreeding, were as likely to be  dichogamous as were self-compatible (SC) plants (Bertin, 1993). This  finding led to a reinterpretation of dichogamy as a more general  mechanism for reducing the impact of 
pollen-
pistil interference on pollen import and export (reviewed in Lloyd & Webb, 1986; Barrett, 2002).  Unlike the inbreeding-avoidance hypothesis, which focused on female  function, this interference-avoidance hypothesis considers both gender  functions.
In many hermaphroditic species, the close physical proximity of 
anthers and 
stigma makes interference unavoidable, either within a 
flower or between flowers on an 
inflorescence.  Within-flower interference, which occurs when either the pistil  interrupts pollen removal or the anthers prevent pollen deposition, can  result in autonomous or facilitated self-pollination (Lloyd & Webb,  1986; Lloyd & Schoen, 1992). Between-flower interference results  from similar mechanisms, except that the interfering structures occur on  different flowers within the same inflorescence and it requires 
pollinator activity. This results in 
geitonogamous  pollination, the transfer of pollen between flowers of the same  individual (Lloyd & Schoen, 1992; de Jong et al., 1993). In contrast  to within-flower interference, geitonogamy necessarily involves the  same processes as outcrossing: pollinator attraction, reward  provisioning, and pollen removal. Therefore, between-flower interference  not only carries the cost of self-fertilization (
inbreeding depression;  Charlesworth & Charlesworth, 1987; Husband & Schemske, 1996),  but also reduces the amount of pollen available for export (so-called  "pollen discounting"; Harder & Wilson, 1998]). Because pollen  discounting diminishes outcross siring success, interference avoidance  may be an important evolutionary force in floral biology (Harder &  Barrett, 1995, 1996; Harder & Wilson, 1998; Barrett, 2002).
Dichogamy may reduce between-flower interference by minimizing  the temporal overlap between stigma and anthers within an inflorescence.  Large inflorescences attract more pollinators, potentially enhancing  reproductive success by increasing pollen import and export (Schemske,  1980; Queller, 1983; Bell, 1985; Geber, 1985; Schmid-Hempel & Speiser, 1988; Klinkhamer & de Jong, 1990).  However, large inflorescences also increase the opportunities for both  geitonogamy and pollen discounting, so that the opportunity for  between-flower interference increases with inflorescence size (Harder  & Barrett, 1996). Consequently, the evolution of floral display size  may represent a compromise between maximizing pollinator visitation and  minimizing geitonogamy and pollen discounting (Klinkhamer & de  Jong, 1993; Barrett et al., 1994; Holsinger, 1996; Snow et al., 1996).
Protandry may be particularly relevant to this compromise,  because it often results in an inflorescence structure with female phase  flowers positioned below male phase flowers (Bertin & Newman,  1993). Given the tendency of many insect pollinators to forage upwards  through inflorescences (Galen & Plowright, 1988), protandry may  enhance pollen export by reducing between-flower interference (Darwin,  1862; Harder et al., 2000). Furthermore, this enhanced pollen export  should increase as floral display size increases, because between-flower  interference should increase with floral display size. These effects of  protandry on between-flower interference may decouple the benefits of  large inflorescences from the consequences of geitonogamy and pollen  discounting. Such a decoupling would provide a significant reproductive  advantage through increased pollinator visitation and siring success.
Harder et al. (2000) demonstrated experimentally that dichogamy  both reduced rates of self-fertilization and enhanced outcross siring  success through reductions in geitonogamy and pollen discounting,  respectively. Routley & Husband (2003) examined the influence of  inflorescence size on this siring advantage and found a bimodal  distribution with increased siring success with both small and large  display sizes.
The length of stigmatic receptivity plays a key role in regulating  the isolation of the male and female stages in dichogamous plants, and  stigmatic receptivity can be influenced by both temperature and  humidity.
[14]  Another study by Jersakova and Johnson, studied the effects of  protandry on the pollination process of the moth pollinated orchid,  ‘’Satyrium longicauda’’. They discovered that protandry tended to reduce  the absolute levels of self-pollination and suggest that the evolution  of protandry could be driven by the consequences of the pollination  process for male mating success.
[15]  Another study that indicated that dichogamy might increase male  pollination success was the study performed by Dai and Galloway.
[16]