Kendall Horan

FISH 350
Article Review #1
13 October 2015
Hagfish Article Review
Between the Lower Silurian and the Upper Devonian eras, for approximately eighty
million years, abundant jawless fishes coexisted with the earths first jawed vertebrates. Deemed
agnathans and gnathostomes, respectively, these two groups of fishes have today switched
places. Since the age in which they began, gnathostomes have grown to dominate the fish world
in number, and agnathans have shrunk to only two successful groups, which make up only 0.2%
of extant craniates. Jawed fish today use their jaws for everything from breeding, to
communicating, to combat which begs the question: how do todays agnathans, the hagfish and
the lampreys, function without jaws? Of what are they capable; what has kept them extant while
so many of their large, armored ancestors have gone extinct? The research put forth in this paper
aimed to broach these questions through a series of trials concerned with comparing the
morphology of the feeding apparatuses of two extant hagfish species (Eptatretus stoutii and
Myxine glutinosa), comparing the feeding kinematics of the same two species, calculating the
forces generated by hagfish musculature when feeding, proposing a physical model of the
hagfish feeding mechanism, and, lastly, comparing hagfish feeding performance to that of
gnathostomes in an effort to evaluate the constraints of jawlessness.
In the end, they found that, when normalized, the total length, mean hagfish feeding
apparatus length, basal plate dimensions, and dental plate dimensions were significantly greater
in E. stoutii than in M. glutinosa. In addition, the deep protractor muscles were significantly

different between the species, the mean forces production of E. stouttii clavatus muscle being
significantly larger than that of M. glutinosa. During food consumption, the cranial movements
in E. stoutii were found to be similar to the cranial movements in M. glutinosa, each depressing
and elevating at near-even rates; the hagfish feeding mechanism was found to be conserved in
both species. When compared to gnathostomes, this information revealed that the jawless
hagfish feeding apparatus is not entirely incomparable to those of vertebrate jaws. Gape size
does not hinder agnathans in the way it does gnathostomes, and, though this may appear, at a
glance, to be the case, lacking jaws and a rigid skeletal system does not reduce the amount of
force hagfish are able to produce overall. Where agnathans do suffer is in speed jawed,
vertebral fishes can bite faster than the jawless hagfish. Though gape size and producible force
are fair advantages, rapid jaw movements are key for catching elusive prey, and the lever system
of the jaw makes for a more efficient use of muscular energy. In the end, this data suggests,
generating considerable muscular forces and attaining wide gape angles were present in the
common ancestor to the craniates, not the common ancestor to the gnathostomes.
The authors of this article succeeded in presenting their data, corroborating said data with
outside references, and arguing their claim in a succinct and understandable way. The diagrams
throughout the article were particularly helpful, as was the organization of the relative sections
and the data itself.

Works Cited
Clark, A.J., & Summers, A.P. (2007). Morphology and kinematics of feeding in hagfish: possible
functional advantages of jaws. The Journal of Experimental Biology, 210 (Pt 22), 38973909. Research Gate, PDF.

Kendall Horan
FISH 350
Article Review #2
20 October 2015

Shark Skin
The placoid scales found on a sharks skin are a marked exception to the popular
description of sharks as purely cartilaginous fishes. Placoid scales are in fact small, tooth-like
elements composed of an outer enameloid layer atop a bone-like layer that surrounds a pulpy,
dentine-based cavity. These scales are commonly called denticles, and erupt in complex threedimensional (3D) shapes through the dermis of sharks. This means, though some of the oceans
most streamline predators, sharks are actually very rough to the touch, and possess skin
somewhat oppositional to a traditional definition of streamline.
This disparity in appearance and function led many scientists to investigate what effect
these denticles have on the flow of water over a sharks body. Scientists studied how denticles
are structured and distributed, and how the surface roughness they create may reduce drag during
locomotion and allow for that streamline swimming. These studies led to practical application:
bioengineers used the basic structure of a sharks skin to develop swimsuits that allow humans to
move faster and more effortlessly through the water.
Because a sharks skin bends and flexes when the shark moves through the water, most
early experiments where synthetic denticles were held atop stiff plates and moved mechanically
through the water are questionable in their accuracy and relevance. To combat those
inaccuracies, flexible pieces of real shark skin were used instead. But this is also an imperfect

testing mechanism, as the preservation of the shark skin compromises its natural flexibility. So,
the authors of this paper set out to design, fabricate, and test a 3D printed biomimetic model of a
sharks skin, complete with both the rough, rigid denticles, and the skins flexible capability.
Using multimaterial 3D printing, the researches designed a swatch of synthetic shark skin
where stiff denticles based on a high-resolution micro-CT scan of the skin of a shortfinned mako
(Isurus oxyrinchus) were placed on flexible membranes in a nonrandom, linear-arrayed pattern.
The scientists then printed a smooth model without denticles, and used that as a control. Both
swatches were put through a series of hydrodynamic tests using a robotic flapping device that
allowed the models to be manipulated in terms of a sharks documented movements. The team
found their fabricated, denticle-laden shark skin resulted in a maximum static drag reduction at
slower flow speeds but increased drag at higher speeds, an increase of swimming speed under
some conditions and a reduction of swimming energy for most conditions, and a significantly
enhanced LEV, or leading edge vortex (a universal mechanism enhancing lift). Though these
findings are significant, that the study of fish skin depends critically on the motion program was
the teams most notable conclusion. Minute changes in how the skin moves dramatically
changes the results found in studies of the skins hydrodynamic properties. That said, the design
and development of this 3D printing biomimetic shark skin is a step in the right direction in the
mission to develop an accurate, repeatable test of a sharks sandpaper skin.
Overall, the presentation of the data was clear and complete, though I did find the authors
repeated themselves in several areas, and the paper lacked a concise conclusion.

Wen, Li; Weaver, James C.; Lauder, George V. Biomimetic shark skin: design, fabrication and
hydrodynamic function. Journal of Experimental Biology 2014 217: 1656-1666.

Kendall Horan
FISH 350
Article Review #3
27 October 2015
Polypterus Walking
Nearly four hundred million years ago, the origin of tetrapods saw the origin of terrestrial
locomotion and the evolution of supporting limbs in fish. Today, a group of scientists studies
Polypterus, an extant analogue of stem tetrapods, in an effort to determine how developmental
plasticity affects the terrestrialization of fish. Their results, along with their methodology,
were put forth in this paper aside several helpful charts and diagrams.
After much analysis of Polypterus reared in opposing environmental conditions, the
authors of this paper found a correspondence between the environmentally induced phenotypes
of terrestrialized Polypterus and the ancient anatomical changes in stem tetrapods. The rapid
response of the skeleton and behavior of Polypterus to a terrestrial environment suggests
environmentally induced developmental plasticity may have facilitated the origin of the
terrestrial traits that led to tetrapods. Essentially, this study, done on a microevolutionary scale,
outlined trends fit for integration into the study of major evolutionary transitions.
Polypterus can function as an analogue for stem tetrapods because it exhibits certain
plesiomorphic morphologies that make it comparable to stem tetrapods. Polypterus, a genus of
freshwater fish, has an elongate body form, rhomboid scales, ventrolaterally positioned pectoral
fins, and functional lungs. This fish can also perform tetrapod-like terrestrial locomotion with its
pectoral fins and is capable of surviving on land. Though other potential analogues exist,

Polypterus already displays walking behavior, and could therefore provide data regarding how
obligatory walking influences gait and skeletal structure.
The scientists reared Polypterus in control and treatment groups under aquatic and
terrestrial conditions, respectively. The placement of a predominantly aquatic fish in an
obligatory terrestrial environment allowed the scientists an in depth look at critical performance,
biomechanical, and anatomical differences between the fish raised in the water and the fish
raised on land. Clear comparisons were found in all three areas: the strain on the musculoskeletal system revealed walking is more energetically expensive than swimming; the land-raised
fish developed a more efficient gait through a conditioned training advantage; and the skeletal
structure, namely that of the pectoral girdle, developed differently enough in the land-raised fish
to greatly impact function and performance. These comparisons are indicative of the
evolutionary changes that enabled the rise of the tetrapods.

Standen, Emily M.; Du, Trina Y.; Larsson, Hans C. E. Developmental plasticity and the origin
of tetrapods. Science.513, 54-58 (04 September 2014). doi:10.1038/nature13708. Web.

Kendall Horan
FISH 350
Article Review #4
3 November 2015
Swimming Noses
This paper chronicles a study of sharks, skates and rays or, elasmobranch fishes and
their acute sense of smell. Olfaction in fishes, meaning a fishs sense of smell, functions by
chemical signals transmitted through the water, and is therefore an important long-range tool
when probing an environment for friend, foe, or food. It is widely believed that compared to
other bony fishes, the cartilaginous elasmobranchs possess superior olfactory sensitivities,
which may allow them certain advantages when finding food and navigating through their
environment. This study, performed by Tricia Meredith and Stephen Kajiura, sought to
investigate any correlation between the structure of an elasmobranchs nose and the overall
effectiveness of its sense of smell. Once a correlation was found and compared across five
species of elasmobranch, the team worked to either confirm or disprove the popular rumor that
elasmobranchs possess a sense of smell far stronger than that of other fishes. In the end, they
found the correlation moot, and the rumor effectively disproven.
In looking for a relationship between the morphology of an elasmobranchs nose and the
sensitivity of its sense of smell, two species of shark and three species of skate or ray native to
similar near-shore environments were examined. From each, the scientists extracted olfactory
organs and summed the number of thin, plate-like, permeable sensors or, lamellae within.
These sums were then studied in conjunction with the whole lamellar surface area to develop a
broader understanding of the morphology of an elasmobranchs nose. Though the sharks

possessed more lamellae than the skates and rays, and size was found to impact surface area,
further testing revealed no significant variation between the thresholds of each individuals sense
of smell. When put in a saltwater swimming pool and subjected to the introduction of an array of
odorous amino acids, each species, despite size and variations in nasal structure, responded to the
odors in the same way. Thus, any correlation between the overall structure of an elasmobranchs
nose and its sensitivity to smell was deemed insignificant.
This determined, the scientists looked to press the rumor that sharks possess the most
superior sense of smell among the fishes, and that the noses of the bony, ray-finned fishes the
teleost fishes fail to compete. Though the noses of sharks, skates, and rays were found to be
quite impressive in their ability to detect a grain-of-rice-sized pinch of amino acid dissolved into
a swimming pool, previous studies show the teleost fishes, with far fewer lamellae, are capable
of detecting the same. This means, despite the structural differences in the nose of an
elasmobranch, any advantage to its sense of smell is not enough to earn it a superior rating
when compared to its teleost relatives. It turns out the swimming noses are no more sensitive
than the other schnozzes of the sea.

Meredith, Tricia L.; Kajiura, Stephen M. Olfactory morphology and physiology of

elasmobranchs. Journal of Experimental Biology. (Impact Factor: 2.9). 10/2010; 213(Pt
20):3449-56. DOI: 10.1242/jeb.045849. Web.

Kendall Horan
FISH 350
Article Review #5
11 November 2015
Molly Sex
Neither bird nor bee, Poecilia formosa, a species of freshwater fish commonly called the
Amazon molly, possesses a reproductive quirk. This quirk allows gynogenesis, which means,
though the mollies must mate with a male to trigger the process, none of the males sperm is
actually used in reproduction, and the female molly produces offspring composed entirely of her
own DNA. Under these biological conditions, P. formosa is a unisexual species made up of only
females. This means the Amazon mollies must perform mating practices with the males of a
related, bisexual species to reproduce. The research recorded by this article looks at Poecilia
formosas mating habits with Poecilia latipinna, or the sailfin molly. The teams main objective
was to determine how and why P. formosa can coexist with P. latipinna when, in theory, a
unisexual species would outcompete a neighboring bisexual species.
Because the Amazon mollies produce only female young who are essentially clones of
the mother, on paper it seems the species should be twice as efficient in furthering itself as its
bisexual counterpart. If one species is twice as efficient as a sister species, reason says the
former would overwhelm the latter, and only the more efficient species would remain. This is
not the case among mollies. Upon investigating how P. formosa mates with P. latipinna, the
researchers found a preference among sailfin males that explains, at least in part, why the
Amazon mollies are kept at bay: the males of P. latipinna prefer to mate with the females of their
own species. Among the females of each species examined, significantly fewer P. formosa

females were found inseminated and impregnated by the P. latipinna males than the P. latipinna
females also examined. This finding suggests the males preference serves to regulate
coexistence between these unisexual and bisexual species.
Though the data collected by this experiment and compiled in this article is solid, the
same experiment has yet to be done on a long-term scale. The male preference pattern
discovered was not stable, suggesting said preference is not the only factor sustaining the
coexistence of the species. Further experimentation will be done, and the results should provide
further insight into how a bisexual species manages to hold its own in harmony with a highly
efficient unisexual species.

Riesch, Rdiger; Plath, Martin; Makowicz, Amber M.; Schlupp, Ingo. Behavioural and lifehistory regulation in a unisexual/bisexual mating system: does male mate choice affect
female reproductive life histories? Biological Journal of the Linnean Society. Volume
106, Issue 3, pages 598606, July 2012. Web.