Fish Final

Muscular System
General Information o Very large proportional of fish body is musculature devoted to locomotive muscle 40-60% of the body mass (good source of protein) o Dont need to conserve body weight can afford to put on more muscle mass as the water supports the fish frame Density of water requires a great amount of propulsive force therefore more muscle is needed to drive movement o Muscles are connected to the head/pelvic girdle anteriorly and to the caudal fin at the tail of the fish Sometimes connected to tendons at the tail (tuna) Run from just below the integument to the core of the animal o Muscle mass is divided vertically into myomeres Heavily vascularized Associated with axial skeleton Myomeres separated by thin, collaginous septa (connective tissue) 2 myomeres exist per vertebra, but due to oblique angle of the myomeres, may span 3-12 vertebra One myomere attached anterior, one myomere attached posterior on each vertebra

o Muscular System Vertical septum is a major division, thick myomere Horizontal septum is also a major division Internally, the myomeres connect to these major septa Red and pink muscle lie in a lateral band along the midline Most is poorly vasuclarized white muscle Proportion of red muscle varies according to lifestyle Need to balance ability to rest and maintain balance and also be able to have quick bursts of speed White burst swimming Red continuous swimming at low speeds Pink intermediate muscle 1

Red Muscle o Highly vascularized, capillary infused, used for sustained swimming o High concentrations of oxygen binding pigments High aerobic capacity, needs lots of oxygen Myoglobin is specific for muscle tissue, has lower oxygen capacity but high oxygen affinity than hemoglobin Favors diffusion of oxygen from blood to muscle o Numerous large mitochondria, high oxidative enzyme activity o Abundant lipid and glycogen stores o Prolonged swimming 20s-200min of constant swimming o Sustained swimming indefinite swimming These types of swimming do not involve fatigue Eg: foraging, station holding, schooling, cruising, and migration Station holding maintaining position in counter current Cruising tuna lack a swim bladder, sink unless they swim o By cross section 5-15% of muscle mass in most species Some species 0% while others are +15% Boarfish 0.5% - chondrichthyes Ratfish 0.6% - chondrichthyes Pollock 11% Flounder very little red muscle Thunniform fishes Carry red muscle deep in the body core o White muscle insulates the red muscle o Need a lot of red muscle for constant swimming to meet their active lifestyle Aids stiff body swimming mode Permits conservation of metabolic heat, allows: o Faster muscle contraction o Greater swimming velocities Tuna are endothermic regionally o High temperature allows for muscle to operate optimally even when looking for prey in cold waters Has the same metabolic output Due to countercurrent exchange system to conserve heat Tuna have a narrow peduncle to allow force of muscle to move tail without move the body because the body is stiff o Conserves metabolic heat, faster swimming regardless of external temperatures White Muscle opposite to red muscle o Poor blood supply and lacks myoglobin o Few smaller mitochondria, has enzymes for anaerobic glycolysis

o Contraction is not dependent on oxygen supply like in red muscle o Anaerobic metabolism converts glycogen to lactate Large spikes in lactic acid Very rapid use of glycogen reserves, ~2mins o Useful for burst swimming 20 seconds Rapid movement of short duration and high speed Subdivided into acceleration (5sec) and sprint (15sec) Locomotion Nonswimming o Terrestrial locomotion Elongate fish move on land as they do in water serpentine motion Eg: eels, can spend limited time out of water Directed flipping, body aligned by the sun Eg: killy fish align with the sun and flip back into the water; could leave water to flee predation after being chased to shallow waters Climbing perch can move up to 3m/min using gill covers as an anchor and vigorous rocking of paired fins and rail Eg: Arabis testudineus has a serrated operculum used to grip the ground so the fins can crawl along o Moving on the bottom Characteristic of benthic sluggish fishes Eg: frog fish angler fish with highly modified pectorals that look like digits, used to walk around, not swimming Sea robins lower rays of pectoral fins separate and move independently like little fingers that are used to locate prey o Jumping and gliding Large fish can clear up to 2.5m salmon, basking sharks Flying fish taxi for up to 9m and then can make up to 12 consecutive flights can cover over 400m Use a hypocercal tail to flap and generate lift 70 beats per second, 60km/h Reason to jump: fish flap when they are being caught due to less resistance in air easier to flap and dislodge whatever is on them Locomotion Modes of forward swimming

o Undulation serpentine with more than one wavelengths present Sinusoidal wave like motion o Oscillation rigid wig-wag motion Exclusive (almost) to the caudal fin Eg: tuna concentrated to narrow caudal peduncle o 5 Basic Modes Anguilliform Entire body undulates; only the cephalic region is kept still Body bent into backward moving waves o Amplitude of waves increases towards the tail o Largest amplitude at the tail Results from the sequential contraction of myomeres Performance improved by lateral compression o Due to dorsal fin merging with caudal fin Not associated with high speeds, benthic fish Carangiform Only posterior portion of the body capable of flexure Undulations confined to the posterior half/third No complete body undulation means side forces to not cancel each other out results in sideslipping o Especially seen in subcarangiform Has a rapid increase in oscillatory movement Coupled with reduction in caudal peduncle Recoil movements is minimized in two ways

o Wave amplitude increase rapidly towards caudal region differs from eels which have a gradual increase in wave amplitude o Depth of caudal peduncle is greatly reduced Mass and/or body depth concentrated towards the anterior deeper than thunniform fish Still median fins increase the span Thunniform Opposite to anguilliform Thrust generated exclusively by extremely narrow caudal peduncle and high still caudal fin Vertebra of caudal peduncle form a thin and rigid unit with lateral keels (fused vertebra), peduncular and postpeduncular joint o Lateral keels are thick peduncle when observed dorsally Creates knife like structure, little resistance o Joints ensure the tail is always flat against the water for greater surface area against the water Tail is in fixed position for high force output Peduncular joint is more anterior than postpeduncular joint Muscles form into tendons which pull back and forth while the body remains rigid to maintain a high propulsive force o Great lateral tendon is a huge tendon used to generate large amount of force o Posterior oblique tendon attaches to the red muscle and the terminal vertebra Red muscles contraction tendon is pulled causing the tail to be bent in that direction o Anterior oblique tendons are attached to the vertebra as well

Nitrogen Excretion
Most nitrogenous waste in fish is produced and excreted as ammonia or urea Ammoniotelism ammonia excretion o Animals that excrete nitrogenous waste as ammonia o Includes most fish includes agnathans and most teleosts (gold fish) o 80-90% of nitrogenous waste is excreted as ammonia (rest as urea) o Ammonia Weak base, highly soluble, readily diffusion across epithelia to leave the body Exists as 2 species in solution NH3 + H3O+ NH4+ + H2O 5

95% exists as insoluble NH4+ (ammonium) o Is protonated, not soluble o Cant diffuse across epithelia o Not permeable Toxic at high concentrations (NH3 and NH4+) Increases internal pH Can inhibit enzymes required for energy generation o Krebs cycle proteins are adversely affected Destabilizes proteins NH4+ can substitute for K+ in ion transporters o Including K-Na-ATPase Devastating effect in the CNS o Disrupts electrochemical gradients o Resistance Fish are much more resistant to ammonia build up than terrestrial vertebrates Fish 100-200M up to 1000M Have less advanced CNS therefore more resistant to high [NH3] Humans - 40M up to 80M before paralysis and toxic effects are felt o Production ammonia production is inevitable Produced by two catabolic processes Amino acid catabolism (mainly) o Requires little to no energy o AAs degraded to -ketoacids by transamination with -ketoglutarate o Ketoacids can be used for energy production, gluconeogenesis and lipogenesis o Primary mechanism is transdeamination 1) transamination amino group of any amino acids is transferred to -ketoglutarate to form glutamate 2) deamination amino group is removed from glutamate by glutamate dehydrogenase to form ammonia and -ketoglutarate o 50-70% of ammonia produced by transdeamination occurs in the liver Rest originates in muscles, kidney, gill, and intestine Purine catabolism o Excretion 80% of nitrogenous waste are excreted by the gills with only trace amounts excreted by the kidney as urine 6

Most of human nitrogenous waste is removed via urine Freshwater Excretion

Passive duffusion of NH3 into water transcellularly and paracellularly Low [NH3] in the blood, but less [NH3] in the water NH3 trapping happens in the water as a proton produces impermeable NH4+ to prevent it from coming back into the cell Proton source is CO2 converted to HCO3- and a proton by carbonic anhydrase Saltwater Excretion

Passive duffusion of NH3 into water transcellularly and paracellularly o NH4+ also exist via passive diffusion paracellularly Only possible due to leaky junction Creates problem of maintaining NH4+ gradient

o NH3 trapping can not happen because SW is buffered cant acidify the water o NH4+ gradient is maintained by exploiting its ability to take K role in basolateral K-Na-ATPase to pump NH4+ into the cell o NH4+ is pumped out of the cell by substituting for a proton in a H-Na exchanger (HNE) Ureotelism urea excretion o Animals thatexcrete urea o Include elasmobranches, coelacanths, some teleosts and lungfish o Urea highly soluble Readily diffuses across epithelia depending on species Elasmobranchs have a high cholesterol:lipid membrane Impeded urea diffusion to maintain high internal osmolyte concentrations Less toxic than ammonia o Production two processes Ornithine-urea Cycle OUC Only ureotelic fish are capable of this Only elasmobranches, coelacanths, a few teleosts, and lungfish that survive periods of air exposure or alkaline environments do this OUC genes encode enzymes necessary for the cycle have been lost from the genome of most teleosts High OUC enzyme activity is detected in many teleosts during embryonic stages OUC genes silenced in adults Carbamoyl phosphate synthase (CPSase; unique) converts glutamine to carbamoyl phosphate o Carbamoyl phosphate OUC urea o Requires energy o Occurs primarily in the liver Uricolysis seen in most fish, breakdown uric acid Also done by ammoniotelic fish o Excretion and Retention at the Gill Urea excretion occurs at the gills same in FW and SW

Passive diffusion of urea into water transcellularly (and paracellularly in SW via leaky junctions) o Minimal amount diffusion Active transport out of the gill by a Na dependent secondary active basolateral membrane urea transporter (UT, symporter) Na gradient maintained by a Na-KATPase o High [Na] in the blood and low [Na] in the cell Urea retention also occurs at the gill that retain urea as an osmolyte to increase body osmolarity elasmobranches Also have counteracting solutes TMAO

o Active transport of urea back into the by a Na dependent secondary active UT antiporter to bring urea back into the blood o Na-K-ATPase maintains Na gradient Ureotelism vs. Ammoniotelism o Ammonia takes little energy to produce Urea production is energy expensive o Ammonia requires a large volume of water for excretion since it occurs by diffusion Urea requires 10X less water for excretion o Ureotelism is better suited for air-breathing fish lungfish lives on land for extended periods of time and has limited access to water 9

African Lungfish o Can live for extended periods out of water in dried mucus cocoons relying entirely on aerial respiration o In water = ammoniotelic o On land = ureotelic due to lack to water making it impossible for ammonia to diffuse from the gills Uses OUC to convert ammonia to urea for safer storage of nitrogenous wastes urea is less toxic than ammonia Gulf Toadfish o Normal conditions = ammoniotelic o Stressful conditions (crowding, confinement) = ureotelic Intermittently ureotelic = excretes pulses of urea Keeps living space clean Urea is sensed by predators so intermittent excretion reduces trail o Transition is accompanied by an upregulation of active OUC enzymes in the liver o Cortisol, stress hormone, is believed to be involved in regulating the transcription of enzymes required for OUC o Negligible ammonia excretion during confinement Other Strategies to Defend Against Ammonia Toxicity on Land o Partial Amino Acid Catabolism Eg: Giant Mudskipper Air-breathing ammoniotelic teleost fish that live in mud burrows When exposed to air, doesnt switch to ureotelism (differs from lungfish) Uses partial amino acid catabolism to generate energy and reduce ammonia production Required enzymes: alanine amino transferase and malic enzyme Uses partial Krebs cycle Pyruvate + glutamate (ALT) -ketoglutarate + alanine o Then -ketoglutarate is shuttled back into Krebs o ATP is produced Malic enzyme converts malate (part of Krebs) to pyruvate) o Shuttles malate out of Krebs cycle to produce pyruvate, thereby generatin a cycle to detoxify ammonia and create ATP Allows amino acids to be used as an energy souce while on land without producing toxic ammonia which would be diffiult to excrete due to a lack of external water o Glutamine Synthesis Eg: Four-Eyed Sleeper Air-breathing ammoniotelic teleost fish that can lives in crevices above river mouths 10

Doesnt switch to ureotelism when exposed to air Uses glutamine synthetase to combine ammonia with glutamate to form glutamine storage of NH3 in a safe form Glutamine = safer ammonia storage than urea Cost effective: 2 mol of ATP per NH3 incorporated into glutamine 2.5 mol of ATP per NH3 incorporated into urea

Circulation and Respiration in Fishes

Circulation o Basics Closed circulatory system Blood flows in a continuous circuit Composed of blood vessels and a muscular pump o Heart Generalities Located posterior to the gills Typical fish heart contains: Sinus venosus Atrium Ventricle Contractile conus arteriosus OR elastic bulbus arteriosus Heart morphology differs between fish groups o Myxini (hagfish) Heart

Most primitive among fishlike species Composed of cardiac muscle Blood enters sinus venosus atrium Ventricle pumps blood into the ventral aorta One way valves between 3 chambers prevent backflow Hagfish have three additional hearts Function to keep blood flowing at high pressure Cardinal heart in head, pumps blood to branchial heart Caudal heart in tail o Medial cartilage plate separates the caudal veins

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o Cartilage plate gets bent back and forth thereby propelling blood o Has skeletal muscle Portal heart near liver, provides blood to the liver o Receives blood from the anterior and posterior cardinal veins o Has cardiac muscle o Petromyzon (lamprey) Heart

Blood enters sinus venosus atrium ventricle Ventricle contraction pushes blood into the conus arteriosus (smooth muscle) which has semilunar valves to prevent backflow o Elasmobrach (shark) Heart

Blood enters sinus venosus atrium Atrial contraction pushes blood into the ventricle Ventricular contraction forces blood to the conus arteriosus Conus arteriosus is capable of contraction o Maintains blood flow to the ventral aorta o Also prevents backflow by bringing conal valves together o Bony Fish Heart

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Sinus venosus atrium ventricle Blood is pumped from the ventricle to the elastic bulbus arteriosus Backflow into the ventricle is prevented by bulbar valves Bulbus arteriosus expands when it receives blood and undergoes elastic recoil to discharge it These processes maintain blood flow to the ventral aorta o Lungfish Heart

Functional three chambered heart (2A, 1V) Atrium is partially divided by the pulmonalis fold Right division receives deox blood via sinus venosus Left division receives ox blood via pulmonary vein Mixing does not occur in the atrium or in the ventricle due to another incomplete partition Conus arteriosus twists 270 so endocardial folds form a spiral valve maintains the separation of ox/deox blood and directs different blood to different vessels Circulatory System o Typical system

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After branching from ventral aorta, aortic arches divide to capillary beds within the gills Blood leaving the gills flows through the dorsal aorta Anterior end of dorsal aorta forms the internal carotids to supply the brain with blood Posterior end of dorsal aorta flows posteriorly to bring blood to systemic circulation o Several arteries branch from the dorsal aorta to supply visceral and somatic tissues o Lungfish Circulation

Atrium and ventricle have partial separation so blood doesnt mix Important when switching between air/water Conus arteriosus has a spiral valve Consists of two endocardial folds which become spirals as the conus twists 270 leads blood to different sets of arches Air-breathing Oxygenated blood from the lungs returns to the heart to be pumped to the III and IV gill arches which lack gills o This goes on to system circulation Deoxygenated (venous) blood returning from tissues flows through gill arches V and VI which is then diverted to the lung to be reoxygenated Water-breathing

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Venous blood is pumped through V and VI gill arches and then diverted to the dorsal aorta There is a shunt that directs blood from V and VI to the lung to be oxygenated in air-breathing and directs blood from V and VI to the dorsal aorta to supply the tissues Blood Composition o Cellular component of blood includes nucleated RBCs and WBCs RBCs (erythrocytes) contain oxygen binding molecule Hemoglobin WBCs (leukocytes) include granulocytes, thrombocytes, lymphocytes, and monocytes o Hemopoiesis Blood cell formation occurs at several sites in fish Spleen erythrocyte formation Head kidney various leukocytes and erythrocytes Thymus lymphocyte production o Hemoglobin Low affinity for oxygen Gill epithelium high PO2 many O2 bound to Hb Tissues low PO2 few O2 bound to Hb The above property ensures uptake of oxygen at the gills and unloading of oxygen at the tissues Bohr Effect o In the presence of CO2, the binding affinity of Hb for O2 decreases o Increases offloading of O2 at tissues and excretion of CO2 at gills o O2 affinity of Hb decreases in the presence of CO2 or acidic pH o In active tissue, large amounts of CO2 are produced CO2 + H2O H2CO3 H+ + HCO3 Results in a decrease of pH increase of O2 unloading at the tissue due to the Bohr effect Therefore this is beneficial to tissues as the decrease in pH provides more O2 to the tissues Respiration o Basics Respiration occurs through the gills Composed of dense capillary beds in the branchial region that serve internal respiration gills supported by branchial arches o Gill Anatomy Lampreys

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Gills lie medial to branchial arch, which is external to gills Contain primary and secondary lamellae which undergo counter-current exchange with water Adult lampreys are usually attached to prey o Water can exit through branchial pouch from external environment tidal ventilation Hagfish

Gills lamellae are enclosed in branchial pouches with muscular walls Scrolling of muscular velum along with contractions and relaxations of branchial pouches create a water current Water current flows in through nostril and continues in one direction across the gills After passing gill pouch water exits via efferent gill duct Elasmobranch

Each gill consists of an interbranchial septum with primary lamellae present on each side of this septum Gill rays within the interbranchial septum lend support

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A branchial arch with lamellae present on both sides of the septum is known as a holobranch A hemibranch has lamellae on only one side of the septum Bony Fishes Gills are covered by a bony or cartilaginous operculum Gills are V-shaped and are supported by a branchial arch Gills are comprised of primary lamellae which are subdivided into secondary lamellae o These secondary lamellae undergo countercurrent exchange with the water Lungfish African lungfish have the greatest modification Have no gills at the III and IV gill arch o Gills are present on the II, V, and VI arches Methods of Ventilation o Ram Ventilation

Involves fish swimming with an open mouth Allows water to move in through the mouth and pass over the gills leading to oxygen uptake Found in large active fish like tuna and some sharks o Dual Pump

Involves a buccal pump and an opercular pump working in tandem Pumps work together to drive water in a continuous, unidirectional flow across the gills Process: Buccal cavity and opercular cavities are closed Buccal cavity expands creating a low pressure driving water into the buccal cavity

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Opercular cavity then expands creating a lower pressure so water flows from the buccal cavity to the opercular cavity Mouth closes, operculum opens, and subsequent compression of both buccal and opercular cavities leads to exit of water through the operculum

o Lungfish In water, lungfish employ same dual pump mechanism When breathing in air, use only the buccal pump to move air into and out of the lungs Occurs in two phases: inhalation and exhalation Exhalation

o Spent air from the lungs is transferred to the buccal cavity by buccal cavity expansion (creates low pressure environment) o Mouth opens, buccal cavity compresses which pushes the spent air out of the mouth (or operculum) into the environment Inhalation

o Expansion of the buccal cavity causes inhalation of fresh air from the mouth o Mouth closes and buccal cavity compresses, forcing the air into the lungs o Gas Bladder Single elongated sac located dorsal to the GI tract Can be filled with air via a pneumatic duct that is connected to the GI tract or with gas secreted from the blood

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Is highly vascularized, can be used to supplement respiration as a respiratory gas bladder Internal walls can be subdivided into many partitions to increase surface area for exchange o Cutaneous Respiration Can occur through skin in air or in water Skin must be highly vascularized Eg: Anguilla anguilla (European eel) has cutaneous respiration accounting for 30% of total gas exchange Eg: Monopterus albus (Swamp eel larva) primarily uses cutaneous respiration during its early life

Sensory Systems
Overview o Octavolateralis System 8th cranial nerve Mechanosensory system lateral line system, free standing neuromasts, put organs Ancient system found in all fish and larval amphibians Not present in amniotes Series of mechanoreceptive organs in the skin Electrosensory system characteristic of vertebrates Ancient system found in many gnathostomes jawed vertebrates, proimarily fishes and platypus Absent from living hagfishes and most teleost fish Electroreceptive organs in the skin Actively send out electrical signals to find prey Or passive system sharks dont create electrical current, just receive it Derived from mechanosensory system Auditory/vestibular system Inner ear ear structure found in all vertebrates Living tetrapods have middle and external ears for airborne sound fish lack these, not hydrodynamic Gives balance, linear and angular acceleration, and hearing Mechanosensory System o Functional unit is the neuromast Cupula jelly-like structure enclosing hair cells Movement of cupula bends hair cells, therefore info is transmitted that that water movement is occurring Hair cells project cilia into the cupula Hair cell sensitivity doesnt differ but the change of cupula displacement can provide stronger impulses depends on cupula shape 19

Support cells support the hair cells o Canal Neuromast Large groups of hair cells, large gelatinous cupula Neuromast in a fluid filled canal of lateral line system o Superficial Neuromast Freestanding, skin neuromast Can occur raised or in pits pit organs Larval fish have superficial neuromasts because the lateral line system is not yet developed Lack gills Use cutaneous respiration o Neuromasts are present all over the body Including the head, tail, and trunk Variations exist between species in Number Placement and branching pattern Canal width Number size and placement of pores Inter-animal variation Functional implications are not yet elucidated 50/50 split between pores of lateral line canals and free neuromasts Dogfish and hammerhead sharks have a lot of pit organs Hair cell is the receptor Substentacular cells Have one long kinocilium and a beveled cluster of 15-30 (up to 150) sterocilli Morphologically polarized Afferent neurons transmit info to the CNS Efferent neurons transmit info from the CNS CN innervated more than SN Electrosensory System o Ampullary Organ Perpendicular to skin surface Subdermal, connects to skin surface Typical of cartilaginous fish 1678 discovery ampullae of Lorenzini Belief was for mucus production, temperature, and salinity sensitive to temp and salt but no functional reason why Canal and tube filled with mucopolysaccharide jelly rich in K Jelly has properties of electrical capacitor Isoelectric and transparent to seawater Sensory cells only have a kinocilium, no stereocilia No efferent innervation of the cell

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Detects weak bioelectric current from surrounding seawater Restricted to the cephalic region Arrangement of organs caries Bilaterally symmetrical, concentrated in cephalic region Lateral line system runs along with ampulla of Lorenzini Used for passive electrolocation Detect weak electric currents generated by prey cardiac muscle contraction and opercular movement Hammerhead shark lower detection limits (% orientations): 70% at <0.1V cm-1 40% at <0.001V cm-1 even down to 0.1nV o Behavior passive electrolocation of prey organisms Experiment to determine what the ampulla of Lorenzini actually do, if bioelectric currents exist, and if sharks make use of this Kalmijn used catsharks and flat fish (plaice) to see if the feeding response is due to electric fields Adding in an odor intensifies the feeding Response isnt due to visual cues Fish is placed in a plastic bag with agar covering it agar is electrically transparent but attenuates optical, chemical, and mechanical stimuli There is an inlet and outlet for water flow Shark locates the fish and focuses on the cephalic region of the plaice where electrical stimuli originate Therefore not a visual cue Instead of a fish in the agar chamber, a bait bag is used Sharks focus on the outlet tube where the smell is Fish is placed in agar chamber with a thin layer of cling film Allows for water displacement to see if mechanical stimuli is used for feeding behavior Cling film shuts down electrical conduction Shark ignored the fish altogether evidence for electrical Electrodes that mimic bioelectric signals are buried in the sand and this elicited a robust and specific natural feeding response Even if chopped up fish was placed in the tank as well, the shark would focus on the electrode and ignore the bait Characteristic Neuromast Ampullary Receptor Function Mechanoreception Electroreception Distribution Head, trunk, tail Head - trunk has midventral line for the lateral line system Receptor Cell at Hair cell Modified hair cell Base - kincilium and stereocilli - kinocilium only Peripheral Afferent and Efferent Afferent only

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Stimulation Stimulus Role

Any displacement Orientation, coordination of swimming, escape, distance touch

- only sends information out Direct current and low frequence alternating current Electrolocation, electrodetection

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