text: In response to decreasing dissolved oxygen level in the environment, fish swim up to the surface of the water column and ventilate at the top layer of the water where it contains relatively higher level of dissolved oxygen, a behavior called aquatic surface respiration (ASR). Oxygen diffuses into water from air and therefore the top layer of water in contact with air contains more oxygen. This is true only in stagnant water; in running water all layers are mixed together and oxygen levels are the same throughout the water column. One environment where ASR often takes place is tidepools, particularly at night. Separation from the sea at low tide means that water is not renewed, fish crowding within the pool means that oxygen is quickly depleted, and absence of light at night means that there is no photosynthesis to replenish the oxygen. Examples of tidepool species that perform ASR include the tidepool sculpin, the three-spined stickleback, and the mummichog.
  But ASR is not limited to the intertidal environment. Most tropical and temperate fish species living in stagnant waters engage in ASR during hypoxia. One study looked at 26 species representing eight families of non-air breathing fishes from the North American great plains, and found that all but four of them performed ASR during hypoxia. Another study looked at 24 species of tropical fish common to the pet trade, from tetras to barbs to cichlids, and found that all of them performed ASR. An unusual situation in which ASR is performed is during winter, in lakes covered by ice, at the interface between water and ice or near air bubbles trapped underneath the ice.
  Some species may show morphological adaptations, such as a flat head and an upturned mouth, that allow them to perform ASR without breaking the water surface (which would make them more visible to aerial predators). One example is the mummichog, whose upturned mouth suggests surface feeding, but whose feeding habits are not particularly restricted to the surface. In the tambaqui, a South American species, exposure to hypoxia induces within hours the development of additional blood vessels inside the lower lip, enhancing its ability to take up oxygen during ASR. Swimming upside down may also help fishes perform ASR, as in some upside-down catfish.
  Some species may hold an air bubble within the mouth during ASR. This may assist buoyancy as well as increase the oxygen content of the water passing over the bubble on its way to the gills. Another way to reduce buoyancy costs is to perform ASR on rocks or plants that provide support near the water surface.
  ASR significantly affects survival of fish during severe hypoxia. In the shortfin molly for example,  survival was approximately four times higher in individuals able to perform ASR as compared to fish not allowed to perform ASR during their exposure to extreme hypoxia.
  ASR may be performed more often when the need for oxygen is higher. In the sailfin molly, gestating females (this species is a livebearer) spend about 50% of their time in ASR as compared to only 15% in non-gestating females under the same low levels of oxygen.