Picture inspired by the previous blog entry.

The origin of walking?

We, as tetrapods, are constitute the largest group of animals in the sarcopterygian clade. Sarcopterygians include, as you may know if you have followed this blog, lungfishes, tetrapods and coelacanths. The common denominator for the animals of this clade are their lobed fins that in tetrapods have become the arms and legs. 

In this group lungfishes are considered to be the closest extant relatives of all tetrapods. A common denominator of tetrapods is, as the name implies, the four paired legs used to propel the body around in the environment. This is a bit different from the coelacanths and lungfishes which still have a set of fins. However the base of their fins are more similar to our arms than you might think, as some of the bones present in their paired fins are equivalent to some of the bones in our arms. The feet is however unique to the tetrapods, believed to be a necessity to be successful in moving about on land. 

Recently a paper was published by King et al. (2011) titled "Behavioral evidence for the evolution of walking and bounding before terestriality in sarcopterygian fishes". In their paper they analyzed the movement of a african lungfish (Protopterus annectens) using video recordings. Their very interesting finding was that this particular species of lungfish uses its pelvic fins as "legs" to move about on the substrate. The fins moved in a synchronized gait as you see in the gaits of tetrapods. So the lungfishes can be in a way bipedal when they move about on the bottom. They can even lift up their body with their pelvic fins, which has previously suggested to be hard without the pelvis connected to the spinal cord as all extant tetrapods.

Link to one of their videos: http://www.youtube.com/watch?v=DgqBzCOfOdE

They conclude that bottom living (benthic) sarcopterygian fishes can lift their bodies up from the substrate and possibly create tracks that later have been fossilized and that the origin of the type of gait used by tetrapods has an earlier origin already present in the water living ancestors of tetrapods.

The lungfish and the tetrapod lineage separated, according to molecular data, approximately 428 million years ago (timetree.org) which would mean that the gait that seem to be common between these two lineages is somewhat older than that. The earliest known fossil tracks was produced by a tetrapod with feets in the Eifedian (early middle devonian, about 395 million years ago) period in todays poland (Niedzwiedzki et al. 2010). This mean that feets have their origin earlier than 395 million years ago, worth noting is that the earliest known tetrapod body fossil is 18 million years younger than these tracks. 

Could the gait style of tetrapods and lungfishes be even older or could it even have been reinvented in several lineages (convergent evolution)? After seeing the videos of the lungfish from the paper described above I couldn't stop thinking of several other non-sarcopterygian vertebrates that also have a similar way of moving about on the bottom substrate using their paired fins as "legs" and "feets". One very striking example is the epaulette shark (Hemiscyllium ocellatum) and its close relatives of the benthic bamboo shark group. Just as the modern lungfishes these sharks are benthic and live in shallow water. These sharks have a habit of crawling about on the bottom substrate and in the coral reef structures using their paired fins (pectoral and pelvic fins) in a similar way as a tetrapod (Goto et al. 1999). This would either suggest that the way tetrapods move have a origin way earlier than split between tetrapods and lungfishes, an origin before the bony fish lineage and the chondrichthyes about 462 million years ago (timetree.org). However since most vertebrate species are not using this type of locomotion but rather swim, just as normal fish, its possible that the jawed vertebrate body plan with its paired fins pave the way for evolving this type of gaits in benthic species that need to move efficiently over the bottom substrate, like the tetrapod ancestors, lungfish ancestors and epaulette shark ancestors. This would also make it difficult to say for sure that the gait style observed in lungfish has the same origin as the tetrapod style, even thought it is likely due to the close relationship between these lineages. It is also possible that the gait style of these two lineages evolved separately in both lineages after the divergence and that the way movement is controlled in vertebrates facilitate evolution of these very similar gait styles.  

Link to video of epaulette crawling about on the substrate: http://www.youtube.com/watch?v=nHJWqYw3IIM

References:

King HM, Shubin NH, Coates MI, & Hale ME (2011). Behavioral evidence for the evolution of walking and bounding before terrestriality in sarcopterygian fishes. Proceedings of the National Academy of Sciences of the United States of America PMID: 22160688


Niedźwiedzki G, Szrek P, Narkiewicz K, Narkiewicz M, & Ahlberg PE (2010). Tetrapod trackways from the early Middle Devonian period of Poland. Nature, 463 (7277), 43-8 PMID: 20054388


Tomoaki Goto, Kiyonori Nishida, & Kazuhiro Nakaya (1999). Internal morphology and function of paired fins in the epaulette shark, Hemiscyllium ocellatum Ichthyological Research, 46 (3)

Boldness and reproductive success in zebrafish

Animals in general react in similar ways to stressful stimuli. There are two basic mechanisms in which animals react to stress. some animals react by being aggressive and actively try to attack the thing that is perceived stressful by the individual while other animals react by being passive and show low levels of aggression. The aggressive individuals have a so called proactive coping style while the inactive non-aggressive individuals have a reactive coping style. Proactive animals often are dominant and reactive animals subordinate in a social interaction. The coping style an animal will show can be determined by many factors for example genotype or environmental factors. The differences between proactive and reactive individuals is not only visible in the behavior but also in their neuroendocrinology. In vertebrates stress is regulated through the so called Hypothalamic-Pituitary-Adrenal Axis (HPA-Axis) or as it is called in fish the HPI axis (where I stands for interrenal, which have the same function as the Adrenals in mammals). The regulation of this axis differ between proactive and reactive individuals where proactive individuals show a lower activity of the axis than reactive individuals. 

The zebrafish, which is a commonly used experimental animal, is a fish that lives in shoals of approximately 10-20 individuals. Zebrafish has its native range in India, Myanmar and Bangladesh where it lives in small streams and flooded plains. Being a social fish species they interact between each other a lot. However how the boldness of an individual zebrafish contribute to the social hierarchy in the group had until recently not been investigated. At the day of my birthday this year my former supervisor published a paper with me among others as a co author (Dahlbom et al. 2011) where we investigated if one could predict social status by testing for boldness in individual zebrafish i.e. if boldness correlates with coping style.

To test this we first tested the boldness of individual zebrafish and then put the fish in a tournament where they met conspecifics in two rounds to isolate the extremes in dominant and subordinate individuals (see Dahlbom et al. 2011 for details). What we could see was that individuals that proved to be bold in the initial boldness screen actually more often ended up dominant (proactive) in a encounter with another zebrafish and thus boldness seem to correlate with coping style shown in the social encounter. 

Recently the first paper citing our paper (Ariyomo and Watt, 2011) was published. They further investigated the effects of boldness and aggressiveness in zebrafish but in the area of reproductive success of male zebrafish. Could it be that bold and proactive males had higher reproductive fitness than shy subordinate males? They saw that if a female was paired with a male it didn't make any difference between bold and shy males on the number of laid eggs by the female. However bold and aggressive males succeeded better with fertilizing eggs. Thus, more eggs in the tanks with a bold aggressive male were fertilized than in thanks with a shy non-aggressive male. 

So to conclude bold zebrafish seem to be more likely to become dominant and be proactive than shy zebrafish. Bold zebrafish males also have a higher reproductive success than shy males by being more successful in fertilizing eggs. Then you might ask, why isn't all zebrafish bold and proactive, why is there still reactive shy individuals? This could be explained by the fact that the different coping styles actually both have their benefits, however in different environments, and thus are both types maintained in the population. 

References

Dahlbom SJ, Lagman D, Lundstedt-Enkel K, Sundström LF, & Winberg S (2011). Boldness predicts social status in zebrafish (Danio rerio). PloS one, 6 (8) PMID: 21858168


Ariyomo, T., & Watt, P. (2011). The effect of variation in boldness and aggressiveness on the reproductive success of zebrafish Animal Behaviour DOI: 10.1016/j.anbehav.2011.10.004

Animal of the week: The yeti crab

"...imagine theres no sun..."

This is probably something John Lennon would have written in the lyrics of imagine if he were a Yeti crab. Fortunately enough he wasn't a yeti crab, because if he was e would probably not have had the cognitive skills needed to write such master pieces. 

The Yeti crabs are (not very skilled in terrestrial music culture), as their name implies, white and hairy. They live in deep water with the first species discovered off the coast of Easter island in 2005 (Macpherson et al. 2005) and the second species the following year (Thurber et al. 2011). The two species have been assigned a genus, Kiwa. 

The interesting thing about these crabs is not that they are white and hairy, although thats pretty awesome in a way, but that they actually culture bacteria on their hairs and use these as their main food source. But the coolness about these critters does not even stop here! The bacteria they culture produces energy from the inorganic gases on the sea floor were these crabs roam about. This mean that the crabs actually live a Sun isolated life fueled purely by the gases of the deep sea floor!  

Some references: 

Macpherson, E., Jones, W. & Segonzac, M. Zoosystema 27, 709–723 (2005). 

Thurber, A. R., Jones, W. J. & Schnabel, K. PLoS ONE. 6, e26243 (2011).

Fresh water stingray facts part III: Thinking rays.

Now you all might think that I shall rename the blog to stingray addict but this will be the last post about rays for a while :).

Rays and sharks in general is often perceived by aquarists as very smart creatures. Compared to many fishes sharks and rays have relatively large brains and in some studies, however very few, been shown to be rather smart. A large part of the brain of sharks and rays is dedicated to processing data from the Ampullae of Lorenzini which is the electro-sensory organ to locate prey. Freshwater stingrays of the family Potamotrygonidae generally have fewer of these ampullae which mean that their brains are not as large as in sea living stingrays. However their brains are still relatively large. The question is how smart are these rays really?

Tool use have been observed in many lineages of the animal kingdom, such as in primates (both human and non-human), in birds and in cephalopods to give some examples. However observations of this kind have been rare in the largest group of vertebrates, namely teleost fish, until now. In a few observations members of the wrasse  family (Labridae) have been seen using stones on the sea floor as anvils while trying to crush clams. This have been observed in three different genera of wrasses so it seem to be a common method used by these fish. The first observation of tool use in wrasses was described in 1995 for the yellow head wrasse.

So tool use have been observed in several species throughout the animal kingdom, and particularly in the bony fish lineage including tetrapods. The cartilaginous fish lineage on the other hand is less studied when it comes to tool use and memory studies. In 2005 a research group used five captive breed Potamotrygon castexi stingrays of the same litter to study cognitive abilities of these rays. They used a plastic tube with one white and one black painted side. On one side of the tube there was a mesh on the inside and the other side was open. Inside the tube they placed a piece of irresistible food for a stingray. The food was, due to the mesh, not accessible from one of the sides of the tube. What they observed was that the rays learned quite well that if they did the right thing they would get the food, they also learned the color on the side of the tube that was associated with the possibility to access food. The way the rays accessed the food was through blowing a jet of water into the tube to sort of flush the food out. This method, by blowing a jet of water, is used by many ray species to uncover pray hidden in the sediment. 

The authors of this paper suggest that this is an observation of tool use in a cartilaginous fish species, using water as a tool to access the food. However if this really is tool use is a matter of definition since its a behavior the rays are born with knowing what to do. One can however argue that in the setting with the test pipe it is in a way tool use since they need to learn to get the food out using the water. I would rather state that the paper is a good study on learning and memory in stingrays rather than an observation on tool use. In any case freshwater stingrays are very interesting and more studies on their cognition would be interesting.

Kuba MJ, Byrne RA, & Burghardt GM (2010). A new method for studying problem solving and tool use in stingrays (Potamotrygon castexi). Animal cognition, 13 (3), 507-13 PMID: 20020169

What is a fish anyway?

One discussion that frequently appear around the lunch table at work is wether humans and other tetrapods are fish. New Oxford American Dictionary define the word fish like this:

"...a limbless cold-blooded vertebrate animal with gills and fins and living wholly in water..." 

Clearly humans and other tetrapods in the majority of the cases lack these attributes. However in many cases the boundaries are not as clear. For a couple of examples we have the lobe finned fish, that possess very limb like fins with even many of the bones of tetrapod limbs; we have the axolotl, a neotenic salamander, that carry gills all there life; we have the mudskippers, that live a substantial time of their life on land and as a final example we have many of the large pelagic shark and game fishes that in a sense are warm-blooded (endotherm). This highlights the difficulties in defining whats a fish and that the definition of a fish, which includes sharks, rays and chimeras as well, may be a old and outdated way of defining this diverse group of animals.

So in some cases the morphological attributes cause confusion, but what groups of animals are included under the definition of the word fish then. Considered, as fish, are cartilaginous fish, ray-finned fish, lobe-finned fish (excluding tetrapods) and the lampreys. However in a biological and evolutionary perspective (figure 1) tetrapods are descendants of lobe-finned fish (the reason for our resemblance within the bone set up of the limbs). So in a biological perspective it is silly to exclude tetrapods from the group of fish, a lungfish is in fact more closely related to you than to a common perch (Tip: http://timetree.org/). And to go further we are, together with the lobe-finned fish, more closely related to the common perch (a ray finned fish) than what cartilaginous fish are to the perch. Still the sharks are considered fish while we aren’t.

In a matter of fact fish in a biological sense fish as a group would include basically all vertebrate species.

Figure 1. Rough tree over the vertebrates. Groups of vertebrates qualifying as fish are marked by the pale red field. Tetrapods are excluded. 

River monsters and a really old citation...

Airing on Animal planet is a show featuring Jeremy Wade called River Monsters. Jeremy Wade is an angler and biologist traveling all around the world fishing for what they call river monsters. A common denominator for these fish is that they are often large and look weird. Some of them are venomous and have caused rare deaths. Often the species he targets isn't very new to science and often common in the aquarium trade. However the show never seem to acknowledge the knowledge present on these species in the scientific literature or in the aquarium hobby. They present each species as if it was new and almost unknown, probably due to dramaturgic effects...

In the episode 'Silent Assassin' (S03E05) we follow Jeremy to the Parana river in Argentina where he follow up stories and victim accounts of stingray "attacks" in the river. According to the locals theres a mysterious stingray living in the river attacking and killing locals and their animals. Instead of doing research in the field of stingrays he sets out on a mission to catch this 'river monster'. Most of the people a bit interested in Elasmobranchs would be aware of the family of stingrays inhabiting the river systems of South America, Potamotrygonidae. However the episode overlook this fact and attribute the attacks to a mysterious ray. This mysterious ray however is a rather well known member of the Potamotrygon genus, actually one of the bigger species if not even the biggest. The species of freshwater stingray in this case is Potamotrygon brachyura. This species was described by Albert Günther (back then named Trygon brachyurus) in the article "A contribution to the knowledge of the fish fauna of the Rio de la Plata" in 1880 so it is not totally new to science. In the episode they wrongly assign this species into the family Dasyatidae and shows a picture of a general whiptail stingray which is the common name of the family. However the Potamotrygon genus is one of four genuses in the family Potamotrygonidae or river stingrays. They are thought this mistake not completely wrong due to the closest extant sea living relatives of river stingrays are thought to be stingrays of the genus Himantura in the Dasyatidae family. 

River stingrays are generally docile and generally only sting victims in self defense. Thats what their poisonous spine are for, self defense, and nothing else. So I think its a overstatement to talk about attacks when often the rays are being stepped on.

Reference
Albert C L G Günther (1880). A contribution to the knowledge of the fish fauna of the Rio de la Plata Annals and Magazine of Natural History, 6, 7-13