A thermal classification system that avoids such irrational results is one based on the

source of heat that predominantly determines the temperature of the animal. Ectothermsare
animals whose body temperatures are determined primarily by external sources of heat.
Endothermsregulate their body temperatures by producing heat metabolically or by using
active mechanisms of heat loss.
Active fishes can produce substantial amounts of metabolicheat, but they have
difficulty retaining any of that heat. Blood pumped from the heart goes directly to the gills,
where it comes very close to the surrounding water to exchange respiratory gases. So any
heat that the blood picks up from metabolically active muscles is lost to the surrounding
water as it flows through the gills. It is thus surprising that some large, rapidly swimming
fishes, such as bluefin tuna and great white sharks, can maintain temperature differences as
great as 10 to 15C between their bodies and the surrounding water. The heat comes from
their powerful swimming muscles, and the ability of these hot fishes to conserve that heat
is based on the remarkable arrangements of their blood vessels.
In the usual (cold) fish circulatory system, oxygenated blood from the gills collects
in a large dorsal vessel, the aorta, which travels through the center of the fish, distributing
blood to all organs and muscles. Hot fishes have a smaller central dorsal aorta, and most of
their oxygenated blood is transported in large vessels just under the skin. The cold blood from
the gills is thus kept close to the surface of the fish. Smaller vessels transporting this cold
blood into the muscle mass run parallel to vessels transporting warm blood from the muscle
mass back toward the heart. Since the vessels carrying the cold blood into the muscle are in
close contact with the vessels carrying warm blood away, heat flows from the warm to the
cold blood by conduction and is therefore retained in the muscle mass.
Because heat is exchanged between blood vessels carrying blood in opposite
directions, this adaptation is called a countercurrent heat exchanger. It keeps the heat within
the muscle mass, enabling these fishes to have an internal body temperature considerably
higher than the water temperature. Why is it advantageous for the fish to be warm? Each
10C rise in muscle temperature increases the fishs sustainable power output almost
threefold, giving it a faster foraging capability!


Sadava, David., Hillis, David M., Heller, H. Craigh and Berenbaum, M. 2011. LIFE:The
Science of Biology, Ninth Edition. U.S.A : Sinauer Associates, Inc.

Tuna and other fast ray-finned fishes can swim at speeds up to 80km/hr (SO mph). Sharks,
penguins, dolphins, and seals are also fast swimmers. As apparent in the examples in
Figure40.2, such animals share a stream lined body contour: a shape that is fusiform,
meaning tapered on both ends. The similar shape found in these speedy fishes, birds, and
mammals is an example of convergent evolution (see Chapter22). Natural selection often
shapes similar adaptations when diverse organisms face the same environmental challenge,
such as the resistanc eof water to fast travel.
Physical laws also influence animal body plans with regard to maximum size. As body
dimensions increase, thicker skeletons are required to maintain adequate strength. This
limitation affects internal skeletons, such as those of vertebrates, as well as external
skeletons, such as those of insects and other arthropods.In addition, asbodies increase in
size,the muscles required for locomotion must represent an ever-larger fraction of the total
body mass. At some point, mobility becomes limited. By considering the fraction of body
mass in leg muscles and the effective force such muscles generate, scientists can estimate
maximum running speed for a wide range of body plans. Such calculations indicate that the
dinosaur Tyrannosaurus rex, over 6 m tall, probably could reach speeds of 30 km/hr (19
mph), as fast as a professional soccer player but not quite the thunderous sprint depicted in
the movie Jurassic
Park.















Certain sharks, bony fishes, and insects also use counter current heat exchange. Although
most sharks and fishes are temperature conformers, counter current heat exchangers are
found in some large, powerful swimmers, including great white sharks, blue fin tuna, and
sword fish. By keeping the main swimming muscles several degrees warmer than tissues near
the animal's surface, this adaptation enables the vigorous, sustained activity that is character
is ticoft hese animals.


Most changes in water temperature as a result of land use activity generally trend upward.
An exception is the release of cold bottom water from stratified artificial impoundments that
may alter the flora and fauna for many miles downstream from a dam. Most other activities
generally raise the temperature of receiving waters with the following effects:
(a) Higher temperatures diminish the solubility of dissolved oxygen and thus decrease the
availability of this essential gas.
(b) Elevated temperatures increase the metabolism, respiration and oxygen demand of fish
and other aquatic life, approximately doubling the respiration for a 10 C. rise in
temperature. Hence the demand for oxygen is increased under conditions where oxygen
supply is lowered.
(c) The solubility of many toxic substances is increased as well as intensified as the
temperature rises.
(d) Higher temperatures militate against desirable fish life by favoring the growth of sewage
fungus and the putrification of sludge deposits, and finally
(e) even with adequate dissolved oxygen, there is a maximum temperature that each species
of fish or other organism can tolerate. Higher temperatures produce death. The maximum
temperatures that adult fish can tolerate vary with the species of fish, prior
acclimatization, oxygen availability and the synergistic effects of other pollutants.
Behavioral thermoregulation occurs when a fish actively seeks out areas of water with higher
or lower temperature. For instance juveniles of the Bear Lake Sculpin (Cottus extensus)
live at the bottom of lakes feeding on the lake floor during the day. Here the temperature
is normally around 5C, however after dark they stop feeding and rise to he surface
where they hang around digesting their food. The temperature near the surface of the
lake is about 14C. By doing this they are able to digest much more food during the night
than they could have done by staying on the lake bottom and so they can grow more
quickly. The difference is considerable, at 5C it would take them 33 hours, or a day and
a half to digest the food they can eat in one day, but at 14C it takes them only 4.5 hours
to digest the same amount of food, so by coming to the surface at night they are making
much more efficient use of their time. And then by returning to the colder water as soon
as its food is digested it slows down its metabolism again and thus conserves energy.

Sometimes the strategies that evolve in simple living creatures are truly amazing and almost
seem to reflect intelligence. Another example of behavioural thermoregulation is the
Sockeye Salmon (Oncorhynchus nerka). Sockeye Salmon are crepuscular feeders,
meaning they hunt for food, and therefore feed only at dawn and dusk. During the full
day and the full night they rest. Scientists have learned that in Canada, in Summer, when
the days are long and the nights are short that the salmon adjust their behaviour to
maximize their efficiency. After feeding at dusk the salmon rest throughout the night at
depths of around 11 metres where the temperature is about 15C, however after their
dawn feeding they retreat to depths of about 37 metres where the temperature is only
about 5C. Why? Because the night is short they need to be warm to digest all they have
eaten before the next feeding session, but because the days are long they have more time
and so can digest their food fully at a lower temperature, and in doing so they reduce
their metabolic costs.

Behavioural thermoregulation is not uncommon in fish, or in reptiles, amphibians and insects.
It is easy to implement and has the advantage of not being permanent, meaning the
animal can, in the best environmental circumstances choose to be warm or cool as it
needs. However it has the problem that the environment is not always at its best, and
even when it is it takes time to warm up enough to allow quick muscular action. Animals
that practice physiological thermoregulation have the edge when the ambient temperature
is not optimum, which even in the sea is quite often.

Physiological thermoregulation is where the fish controls its core body temperature by
means of internal physiological and metabolic activities. This is also how we maintain our
body temperatures, but while it is universal among mammals it is rare in fish. It occurs in
only a few species, all of which are marine and swim constantly. These include various Tuna,
some Mackerels, the Mackerel Sharks (Lamna nasus and L. ditropis) and the Great White
Shark (Carcharodon carcharias).
Both sharks and bony fish that maintain an increased body temperature do so by means of
a counter-current exchange system whereby blood vessels carrying blood that is hot as a
result of muscular activity pass along side, and give up some of their heat to, blood that is
going to parts of the body the animal wishes to keep warm. The organ where the heat
exchange takes place is called retia mirabilia. A fish may have more than one retia mirabilia,
Mackeral Sharks for instance have three, one in the swimming muscles, one in the body
cavity near the guts and one around the brain.
As with any other successful adaptation there are many other minor changes that work
along side the major one in harmonizing the animals metabolism. For example Albicore Tuna
have evolved a unique form of haemoglobin that increases its affinity for O2 as it gets
warmer while its CO2 affinity increases as it gets cooler. This is the reverse of the normal
situation, but it does prevent O2 from being lost from the warming arterial to the cooling
renal blood in the retia mirabilia.
The next step up from recycling the heat from muscular effort to heat parts of the body,
which obviously only works while the animal is using it muscles (which is why all the
species that do so are constant swimmers), is to use purely metabolic processes to generate
heat. The only example I know of in the fish is the Swordfish (Xiphia gladius). Swordfish
have specifically modified eye muscles with an exceptionally high mitochondrial volume.
Mitochondria are the organelles wherein food is turned into energy, they are found in every
cell of your body. The swordfish does not use these special eye muscles for anything except
normal eye movements (which does not generate much heat) and to heat the blood going to
its brain. In other words the mitochondria work purely to make heat for the purpose of
keeping the precious brain warm, it is not merely recycling heat that was a byproduct of
normal muscular activity as are the sharks and the tuna. This is much closer to the
mechanisms used by birds and mammals to maintain their body temperatures. Evolution
takes a long time in comparison with a short life spans, but who can say what fish will be
swimming in the sea and oceans in another 50 million years.
Remember, as I said above chemical reactions occur more quickly at warmer
temperatures, warmer guts digest food more quickly, a warmer heart beats more strongly,
warmer muscles contract more strongly and respond more quickly and warmer brains work
better. The contraction power of the muscles of Bonito (Sarda chiliensis lineolata) doubles
with a 10C increase in temperature. All this can give a predator and advantage that makes
the difference between life and death. All the fish and sharks that maintain a raised body
temperature are active constant hunters. The tuna and mackerels hunt smaller fish and the
sharks hunt the tuna and mackerel amongst other things. Endothermic animals need to eat a
lot more than ectothermic ones, sometimes as much as 3 or 4 times as much.
Beberapa jenis ikan seperti ikan hiu dan tua telah memiliki kemampuan untuk
mempertahankan adanya perbedaan suhu antara suatu bagian tubuh dengan bagian tubuh
yang lain. Ikan tunda juga mampu meningkatkan laju reaksi metabolik di tubuhnya, terutama
pada otot yang digunakan untuk berenang dan pad asaluran pencernaannya sehingga bagian
tersebut selalu lebih panas daripada bagian lainnya.kemampuan tersebut dimiliki karena
adanya heat exchanger (penukaran panas) pada tubuhnya. Heat exchanger bekerja dengan
prinsip counter current (arus bolak-balik). Selama heat exchanger bekerja, darah pada
pembuluh arteri yang lebih dingin mengalir dari insang berdampingan dengan pembuluh vena
yang suhunya lebih tinggi, yang mengalir ke insang . dengan cara itu, panas dapat
dipindahkan dari darah vena ke darah arteri, dan masuk kembali ke dalam organ segingga
suhu pada otot renang tetap berkisar antara 12-15oC, lebih tinggi daripada suhu air.
Berbeda dengan lingkungan akuatik, suhu di lingkungan teestrial selalu berubah denga
variasi yang cukup besar. Perubahan suhu ini sangat mudah kita rasakan, misalnya dengan
membandingkan suhu udara pada siang dan malam hari, pada hari yang sama di suatu kota.
Cara terpenting yang dilakukan oleh hewan ektotermik terstrial untuk memperoleh panas
ialah dengan menyerap panas/radiasi matahari. Hewan dapat meningkatkan penyerapan panas
mayahari dengan cara mengubah warna permukaan tubuhnya dan menghadapkan tubuhnya
ke arah mayahari. Invertebrata ektotermik terstrial dapat mengubah warna tubuhnya menjadi
lebih gelap untuk memperoleh panas matahari lebih banyak. Hewan yang melakukan cara ini
anyara alin belalalang rumput (belalang hijau) dan kumbang.
Categories: Anatomy, Fish, Locomotion, Physiology
Thunniform swimming depends on a large, lunate tail that is joined to the rest of the
body via a narrow peduncle. Whilst the tail flicks backwards and forwards, so propelling the
animal, the rest of the body hardly moves sideways. It comes as no surprise to see that large
marine animals that swim fast, typically range very widely (sometimes for thousands of
kilometres) and can often dive to considerable depths, typically have a fusiform body, a
prominent tail and stabilizing fins. Familiar examples include many marine mammals
including dolphins, the extinct ichthyosaurs, the sharks and tuna. In the last two cases,
however, there is a very striking convergence that, as Jeanine Donley and colleagues rightly
emphasize, is much more than skin deep. In the case of the sharks the particular comparison
rests on the lamnids, although a similar case applies also to the somewhat less well-known
alopiid sharks.
Thunniform swimming in lamnid sharks and tuna
Both tuna and shark progress by what is known as thunniform swimming. In essence this
depends on a large, lunate tail that is joined to the rest of the body via a narrow peduncle.
Whilst the tail flicks backwards and forwards, so propelling the animal, the rest of the body
hardly moves sideways. Grey reef shark This is in marked contrast to most fish which
locomote by throwing the body into a series of sinous waves, generally referred to as
carangiform and is at its most pronounced in the angulliform locomotion of the eels. In all
cases, of course, the locomotory power is provided by the muscular contractions, but in
thunniforms the power is transmitted to the tail in a very specific fashion.
It is necessary to know that in fish there are typically two types of muscle. Usual
locomotion is achieved by contraction of aerobic red muscle that is usually located on the
body margins, and whose contractions can throw the body into sinous waves. The white
muscle is only used for power bursts, and because it can function without oxygen it is
referred to as anaerobic. Understandably power bursts can only be of short duration because
the oxygen debt has to be paid off (a direct analogy is the build-up of lactic acid in our calf
muscles when we run too fast for the legs to be supplied with enough oxygen). In lamnid
sharks and tuna, the fish have independently arrived at an almost identical solution to achieve
thunniform locomotion. The main mass of red muscle is moved more centrally, and becomes
largely independent of the rest of the muscle blocks so when it contracts it slides semi-
independently. Second, and critically, the power of the muscular contractions is transmitted
to the tail via a system of tendons. Crucially, although the function of the tendons is
effectively identical, their origin in lamnid sharks is different from that of tuna. Thus, as ever,
there is an evolutionary footprint.
Thermoregulation in sharks and tuna
Another key innovation of this system is the independent evolution of thermoregulation
and endothermy ("warm-bloodedness"). Endothermy has evolved independently many times
e.g. in other vertebrates such as birds and mammals, and also many groups of insects, such as
the sphingid moths. Bluefin tuna The principle area of heat generation in endothermic fish is
in the alimentary canal, but heat is generated in different regions in the shark and tuna.
Secondary areas of heat generation are also found in the brain and eye regions of sharks and
tuna, and it is possible that this assists visual acuity and cognition, at least in the shark.
One critical anatomical feature in the endothermy of shark and tuna is the evolution of a
counter-current blood system. Typically, this consists of a network of blood vessels (rete
mirabile, or wonderful network) whereby blood flowing in either direction can exchange
heat (or in cases such as the fish swim bladder, respiratory gases). In effect, heat that is being
transported out of the zone of thermal generation is constantly transported back so as to
maintain a core temperature higher than other areas of the body. This has a potentially
disastrous consequence because the solubility of oxygen decreases with increasing
temperature, so the warmth that is being returned to the core region by the blood flow ought
to contain less oxygen, a hardly helpful development given that this is a region of high
metabolic activity. The trick to solve this physiological problem has been to induce a so-
called reverse temperature effect in the haemoglobin (the protein that carries the oxygen), and
unsurprisingly this has evolved independently in shark and tuna.

All animals face a dilemma. Optimal efficiency in terms of metabolism, digestions,
strength and movement is improved as the body temperature increases. However, the higher
the internal temperature, the more energy is spent maintaining it.
Sharks in general are poikilothermic or, informally, cold-blooded. This means that the
internal temperature of their body is as cold as the waters in which they live. If the
temperature around them drops, so does their body temperature.
Therefore, these animals are able to conserve energy by rising to the top of the ocean,
where the water is warmer, so that their body temperature rises without much extra effort
from their side. In so doing, digestion is aided, muscle activity is improved and metabolism is
boosted. In an attempt to conserve energy, they will sink to the colder depths, where all of
their body functions slow down. The colder the waters, the more energy the shark uses in
trying to hunt, digest food and travel.
There are some sharks that are homeothermic, which implies that their body temperature is
always slightly higher than that of the water of their environment. This extra heat is generated
by a strip of red muscle (as opposed to white muscle) that runs along the centre core of the
fishs body. This aerobic muscle is surrounded by the rete mirabile, a web of veins and
arteries around the muscle that uses a countercurrent flow of blood within its complex to
exchange heat, gases and ions faster and better.
The blood that has been warmed by increased muscle activity is carried through this
network and the heat is exchanged back into the blood travelling to the muscles. Heat is thus
generated and maintained regardless of the temperature of the water outside the body; it is
constant. This is unlike poikilotherms, whose body temperature is dictated largely by the
water in which they are swimming. The Great White Shark and the Mako Shark are common
examples of homeotherms.
Some sharks and other fish are even able to keep certain areas of their body (such as their
eyes and brains) warmer than the others so that their functionality is not negotiated even
when their muscles and metabolism slow down.
Some sharks display behavioural thermoregulation, which refers to their choosing specific
habitats or behaviours based solely on the water temperature. An example of this is rising to
the top during the safer night hours to digest food faster using less energy. The seas
temperature is not constant or all that predictable. Therefore, the sharks that are able to
implement behavioural thermoregulation have an advantage over other poikilotherms that
cannot.
The warmer the body needs to be kept, the more the animal needs to eat. This is to
increase its energy stores. As the body temperature increases, the muscles work stronger and
harder, the heart beats better, the guts digestion improves and the rate of metabolism
increases.
Thermoregulation (Rete mirabile)
Most bony fish (except for tuna) and most sharks are ectothermic. This means that these
animals are unable to control their own body temperature through their metabolism, but that
their body temperature is controlled by the surrounding water temperature. Exceptions among
the sharks are the mackerel sharks (lamnidae, inducing the great white shark and the maco
shark) and probably the common thresher (alopidae). They are endothermic, which means
that they can regulate their own body temperature.
There is a network of blood capillaries (Rete mirabile) between their red swimming
muscles (in a mackerel shark with a length of 1m, it has a surface of 4m!), which acts as a
heat exchanger. The heat created through muscle activity is transported by the blood in the
vessels that takes oxygen-depleted blood to the gills. Blood vessels with thin walls and cold,
oxygen-rich blood from the gills are running in the opposite direction along the heated
vessels and are warmed this way.
Thus, heat is transferred to the blood flowing into the body, while the blood flowing
towards the gills cools off already and cannot lose as much heat to the outer medium
anymore. This principle of opposite streams is so efficient that hardly any body heat is lost
through the gills, and the temperature inside the body is 5 - 14C above the temperature of
the surrounding water. This way, the red muscles can work more actively and particularly
efficiently. They regulate the temperature in fast-swimming sharks perfectly.
Behavioral Thermoregulation
Most species of fish show behavioral temperature regulation and thereby tend to be found
within a specific temperature interval. This behavior is advantageous for the fish since several
processes, such as acid-base balance, metabolism, blood flow, locomotion, ion and water
flux, as well as enzymes and the nervous system are influenced by temperature (Crawshaw
1979). The mechanism behind behavioral thermoregulation is controlled by the central
nervous system and is comparable to similar mechanisms in higher vertebrates. Fish can
register even small temperature changes. As small a difference as 0.03-0.1C has been
reported as whole-body sensitivity (Crawshaw 1979). The temperature interval a fish is
limited to can be subdivided into the resistance, tolerance and preference zones (Figure 1).
Inthe resistance zone, the fish meets extreme temperatures and its survival time is strongly
dependent on the exposure time (Jobling 1994). In the tolerance zone, the fish survives
without temperature-associated problems, while the preference zone, centered on the
preferred temperature, is thought to be the zone where physiological processes are optimized
(Jobling 1994). Physiological processes that have been shown to be optimized at the preferred
temperature are appetite, growth rate, active metabolic rate, the scope between standard and
active metabolic rate, and sustained swimming speed (Figure 2) (Brett 1971, Beitinger and
Fitzpatrick 1979, Jobling 1981).

Figure 1. Temperature polygon showing the different temperature zones. The resistance
zone is between the Critical Thermal Maximum (CTM) and the Upper Incipient Lethal
Temperature (UILT). LILT = Lower Incipient Lethal Temperature. The preferred
temperature, surrounded by a narrow preference zone, can be found at the intersection of the
Acute Preference Line (AP) and the Line of Equality (LE) (from Jobling 1981).
Pengaruh suhu terhadap pertumbuhan ikan
Suhu media berpengaruh terhadap aktifitas enzim pencernaan. Pada proses pencernaan
yang tak sempurna akan dihasilkan banyak feses, sehingga banyak energi yang terbuang.
Tetapi jika aktifitas enzim pencernaan meningkat maka laju pencernaan juga akan semakin
meningkat, sehingga tingkat pengosongan lambung tinggi. Tingkat pengosongan lambung
yang tinggi menyebabkan ikan cepat lapar dan nafsu makannya meningkat. Jika konsumsi
pakan tinggi, nutien yang masuk kedalam tubuh ikan juga tinggi, dengan demikian ikan
memiliki energi yang cukup untuk pertumbuhan.
Suhu media juga berpengaruh terhadap aktifitas enzim yang terlibat proses katabolisme
dan anabolisme. Enzim metabolisme berpengaruh terhadap proses katabolisme
(menghasilkan energi) dan anabolisme (sintesa nutrien menjadi senyawa baru yang
dibutuhkan tubuh). Jika aktifitas enzim metabolisme meningkat maka laju proses
metabolisme akan semakin cepat dan kadar metabolit dalam darah semakin tinggi. Tingginya
kadar metabolit dalam darah menyebabkan ikan cepat lapar dan memiliki nafsu makan tinggi,
sehingga tingkat konsumsi pakan meningkat. Konsumsi pakan yang tinggi akan
meningkatkan jumlah energi yang masuk ke dalam tubuh. Energi ini akan digunakan untuk
proses-proses maintenance dan selanjutnya digunakan untuk pertumbuhan.
Suhu media yang optimum akan mendorong enzim-enzim pencernaan dan metabolisme
untuk bekerja secara efektif. Konsumsi pakan yang tinggi yang disertai dengan proses
pencernaan dan metabolisme yang efektif, akan menghasilkan energi yang optimal untuk
pertumbuhan.
Proses metabolisme ikan umumnya meningkat jika suhu naik hingga dibawah batas yang
mematikan. Berdasarkan hukum vant Hoff, kenaikan suhu sebesar 10C akan menyebabkan
kecepatan reaksi metabolisme meningkat 2-3 kali lipat dibandingkan pada kondisi normal.
Kebutuhan protein pada ikan untuk mendapatkan pertumbuhan yang optimum sangat
dipengaruhi oleh suhu. Contoh pada suhu 20oC pada ikan Channel Catfish (Ictalurus
punctatus) memperlihatkan pertumbuhan optimum dengan kadar protein 35 %, sedangkan
pada suhu 25oC membutuhkan protein 40%.
Pendekatan-pendekatan yang dapat dilakukan untuk memaksimalkan pertumbuhan ikan.
Ectotherms have a capacity to cope with the changes in temperature that occur in the
environment by selecting favorable habitats for using behavioural responses [4]. The
preferred temperature representsa thermal interval in which the processes that control activity
are not affected and their efficiency is therefore increased and optimized [7-9]. Fry [10]
defines the final preferendum as the temperature around which all individuals (of a given
species) can congregate, regardless to previous thermal history before being placed in the
gradient [11,12]. Jobling and Kellog [13,14] have shown that organisms select temperatures
in proportion to quantity of metabolic energy necessary for maximize different biological
processes such as metabolism, swimming speed, growth. Body temperature (Tb) is the most
important ecophysiological variable that affects the function of ectotherms. Virtually all
aspects of behavior and physiology of ectotherms are sensitive to Tb [15, 16]. The
thermoregulatory behaviour is used by ectotherms to reduce the spatial and temporal impact
of environmental temperature under Tb [15]. Thermal preference studies conducted in the
laboratory propose a vital link in the environmental, physiological, and adaptive behavioral
tactics used by aquatic organisms [17].

Fish and other water-breathing animals face a serious heat loss problem because
water has high heat capacity andlow oxygen content. Must move a lot of water over gas-
exchange surfaces (gills) to get O2. Air breathers can move less air over gasexchange surface,
and air has low heat capacity, thus possible to heat inspired air to body temperature. Energy
cost of heating water (large volume, high heat capacity) would be too great water breathers
constrained to poikilothermy (insulation can't solve this problem).
Heterothermic fish. Some fish (notably tuna) have evolved a counter-current heat
exchange mechanism that allows a part of the body (red swimming muscle) to be kept above
ambient temperature. Arterial blood passing through gills is unavoidably cooled.
Heterothermic fishes' circulatory system routes cooled arterial blood (after gills) near
the surface, so core body heat isn't conducted to it (L maximized, in equation for conductive
heat flux). When this blood is routed into red muscle deep in body, it passes throughrete
mirabile, which is a counter-current heat exchanger a system of veins and arteries running
in opposite directions in contact with each other. Cold arterial blood is warmed by passing
venous blood, which has been heated up by metabolic activity in muscle. By passively
moving heat from venous to arterial blood, some of the metabolic heat carried away by veins
is returned to muscles. Consequently, red muscles in tuna's core can be kept up to 15o C
above water temperature.

Temperatures at which fishes lived in the outfall area during summer 1970 were
positively correlated (p < 0.01) with the degree to which they were concentrated in the outfall
area during the two preiious summers. The strength of the correlation argued that, during
summer, temperature was the primary factor governing not only distribution of fishes within
the outfall area, but also the abundance of fishes in the outfail area relative to unheated parts
of the littoral zone.

Behavioral regulation of environment has been an under-emphasized feature in the
ecol- ogy of fishes. Fishes are mobile organisms, able to swim through their habitat at
sustained speeds of a few body lengths per second (Bain- bridge 1960). In aquatic organisms
with lesser potential for voluntary movements, physiolog- ical adaptation is perhaps the
primary strategy for coping with environmental variation. Physiological mechanisms
allow fishes also to successfully occupy diverse habitats and toler- ate temporal fluctuations
of environment. But, if the potentially available environment is heterogeneous, a fish can
behaviorally regu- late the conditions it must experience by spending more time in one
subset of environ- ment and less or no time in other subsets. Temperature heterogeneity of
some aquatic habitats is greatly increased by their reception of heated effluents from steam-
electric plants. Although the fraction of habitat near the point of effluent discharge (outfall
area) may be warmed by several degrees Celsius, tempera- tures throughout most of the
receiving water mass typically are unaltered. For example, only about 0.3% of the water at
a depth of 0.5 m in Lake Monona is warmed more than 1 C by effluent from a power
plant; the per- centage of total volume so warmed is much smaller.
In a habitat partly heated by thermal efflu- ent, then, temperatures experienced by
fishes depend on where they live within the habitat. If fishes were randomly moving
particles, the number exposed to elevated temperatures and the time each individual is
exposed would be small. Fishes typically are not randomly dis- tributed within a habitat;
rather, they tend to
be concentrated in some areas and scarce or absent in others.
Thus, the effects of steam-electric power production on fishes must depend first
on their behavioral responses to elevated tempera- tures and to other features of the outfall
area. It matters little whether in the laboratory yellow perch (Perca fluvexens), for
example, can grow at 30 C or can survive 33 C for 1,000 min if yellow perch never occur in
an outfall area where temperatures exceed 29 C. The more important question is whether
yellow perch given a choice will invade water of a particular temperature and, if so, for how
long and for what reasons, ecologically prudent or otherwise. Only when fish are likely to
occur voluntarily or by necessity under certain con- ditions does it become ecologically
important to know the consequences of that exposure- consequences relative to survival,
growth, and reproduction.