Note for BSC 373 with Professor Harris at UA-Lecture 7
Note for BSC 373 with Professor Harris at UA-Lecture 7
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Date Created: 02/06/15
Properties of Water Water versus Air see Box 41 Density 800x denser than air Bunyancy Viscosity 18X more viscousthan air Oxygen content 209 milliliters 02 in a liter of air 50 ml 02 in a liter orwater or less Heat Capacity and Heat Conductivity Specific heat of Water 340 that of air Conducts heat 24X rasterthan air Electrical conductivity Problems of Breathing in Water Challenge is to extract 02 39om water and distribute it to the cells in the body fast enough to meet metabolic demands and prevent lactic acid buildup No easy task see Box 41 Concentration of oxygen 7 0 air is appruxirnately 2W by vuiurhE 7 0 water lt i byvuiurne Density and viscosity 7 H20 8qu rncire dense than air 7 H20 aux rncire VlSEDUS So more energy is required to just move water across respiratory surfaces sh use 10 ofthe 02 it gets from waterjust to keep breathing muscles going Problems of Breathing in Water Challenge is to extract 02 39om water and distribute it to the cells in the body fast enough to meet metabolic demands and prevent lactic acid buildup No easy task Concentration of oxygen Density and viscosity Solubility 7 Sulubility decreases as temperature increases 7 Sulubility decreases as saitsdidtes increases suitingout effect 7 5d vvarrn water has iess 02 than cciid water and saltwater has iess 02 than freshwater 91012 91012 Buccal and Opercular Pumps Oxygen ow in shes is unidirectional Contrasts with terrestrial vertebrates in which 02 ow is bidirectional in ow and out ow associated with dead air Water is almost always sucked in through the mouth and exits by way of he opercular openings on each side ofthe head Unidirectional ow negates waste associated with deadair space Makes possible the opportunity to maintain a constant gradient for O2 to diffuse from the water into the blood via the gills Buccal and Opercular Pumps mm VIEW a Man mm mam unknown siphon mom mu m Buccal and Opercular Pumps Breathing Cycle Timing ofthe expansion and contraction ofthe buccal and opercular cavities ensures that the pressure in the buccal chamber exceeds that ofthe opercular chamber throughout nearly all ofthe respiratory cycle This creates a nearly steady ow of water from the buccal chamber to the opercular chamber passing over the gill lamellae which have blood owing through them in the opposite direction 91012 Buccal and Opercular Pumps Mam or um um Mann cu car van clone meal marlne39 maturing nmwu manual Q Cocteau yawn coon 5mm wmn chem Pmssua pump Vase Gill Structure and Function Each gill arch bears a number of gill filaments holobranchs each of which is made up of two halves hemibranchs Fine subdivisions on hemibranch are gill lamellae major respiratory portion of gills Total surface area of the gill lamellae averages about 5 cm2 per gram of body weight Gill arch Halnhianzh Capillary m in lamella ramming lamenlal Muir Limellze Gill arch Gill lilamelil Ellmnt lamzlla S lnwillg lilamenlal amnion nl him may Him through mum m gm Illaquotth Blood Circulation and Respiration Deoxygenated blood reaches the gills by way of the afferent branchial arteries Oxygenated blood from the gills passes into the efferent branchial arteries and into the body Blood flows through the lamellae in the opposite direction of water flow across the lamellae countercurrent Blood with higher 02 content meets water with highest 02 content so that 02 diffuses into the blood along the entire length of the lamellae The effect is an extremely efficient interchange of O2 and CO2 between water and blood Suzme imam m an mu m Dmchr cl watt 9ch Emuanx cued maul 91012 Evolutionary Trends in Respiration Agnathans Intake is through the nostril hagfishes Ventilation pump velum 116 gill sacs with countercurrent setup When the hagfish is buried inside prey water comes in and out through the gill opening behind the last gill mi pouch Lampreys expand and contract the branchial area causing water to flow inout PM practical when head buried WW in prey Nwslm em openlng Mucous gland apB UVES A Gm openings Elasmobranchs Water intake is through mouth and spiracle Ventilation can be either ram mouth or pump mouth and spiracle 2 5 individual gill slits vs single emsms operculum Structure Gill arch and ray supports gi I V laments gIII laments Gill rakers protect laments and collect food aorerem branchial artery gill lilamsnts ellerem branchlal artery Teleosts Ventilation ram mandatory in tunas andor pump buccal opercular cavity pumps Structure Surface area of gills is correlated to activity levels Evolutionary Trends in Respiration w Tum Jl CuMMnmw of am Du 5quot Lum mumym pa mm u t 75 l mmutqmmn H mm mm murmur lt H 5 m mam 3pm Amwnnmlmlmm m a 15 n m u y 7 s 5 Mimvli mu my w mum n u lt5 AWNmm WW 5 X n 0 ll 1 m u 7 z w m in I Ariwc 3pm l2 1x u 1 u 7 w 2 1 2 m I mm mm l39cn li m l 2 l rm mmm Hy Amu 7 2 2n a mm c s 1 Oxygen Respiration etc When 02 needs increase What are he options Breathe more often increase ventilation frequency Take bigger gulps of water increase stroke volume Recruit more gill lamellae Lamellae at tips receive less water and blood ow As water ow increases lamellae at tips receive more ow As the lamellae at the tips get more blood ow this generates higher blood pressure and dilation Oxygen Respiration etc Costs of breathing High density and viscosity of water requires energy for ventilation of gills Branchial pump 1015 metabolic energy Humans cost of breathing at rest is 2 ofmetabolic energy Can lower metabolic costs by ram ventilation Requires an active lifestyle Rapidly moving water salmonids Constantly swimming shes tunas and some sharks Tranfers work of ventilation to locomotory muscles Air breathing Accessory airbreathing structures of bony shes are all modi cations ofthe epithelia ofthe alimentary canal gill chamber or diverticula emerging from these parts 91012 91012 Air Breathing All associated structures have in common a specialized epithelium that is often equipped with branched treelike outgrowths to increase surface area Lying underneath this thin epithelial layer is a rich network of capillaries Gills are never used for air breathing because they are poorly adapted to it Gill lamellae are not stiff or rigid collapse when out of water suffocation Watking my Sacnnd gill elm Air Breathing All airbreathing organs function just as absorbers of oxygen no involvement with excrete COZ done by gills andor skin Evolutionary trend toward the separation of organs dealing with 02 uptake and CO2 discharge Biological significance of air breathing Survival in oxygen poor habitats Utilization of terrestrial food resources Abandon drying ponds to search for better habitat Invade new territories and expand distribution Labyrinlhine plates Oxygen Uptake and Release The tendency of blood to take up and release oxygen is described with an oxygen dissociation curve The amount of oxygen taken up by hemoglobin Hb increases with oxygen tension partial pressure The halfsaturation point P50 is defined as the oxygen tension at which blood is halfsaturated The dissociation curve is typically sigmoid in shape this is because of tetrameric molecular structure Oxygen Uptake and Release Change in pH influences affinity of Hb for oxygen Hemoglobin Saturation Blood pH drops where there is higher carbon dioxide areas of metabolic activity Conformation of Hb changes and reduces its affinity for oxygen lower halfsaturation point This is called the Bohr Effect occurs in all animal hemoglobins In some shes there is an additional effect low pH lowers the oxygen capacity of Hb reduction of asymptotic saturation This is called the Root Effect As a result Hb unloads oxygen in tissues where there is metabolic activity Oxygen Uptake and Release Bohr Effect 9quot P Root Effect Lower H 0 Higher pH Lower pH Hemoglobin Saturation Partial Pressure of Oxygen p02 Partial Pressure of Oxygen p02 Oxygen Uptake and Release Diversity among fishes in dissociation curves and Bohr effect Relative to Hb in an active fish such as mackerel the oxygen capacity of Hb in a fish living in lower oxygen waters toad sh is lower and the Bohr effect is lower the toadfish is less active and there isn t as much oxygen unloading at peripheral tissues 7 Hemoglobin Saturation as 16202520354 Partial Pressure of Oxygen p02 91012 91012 Oxygen Uptake and Release Ehmmatton ofCOZ e h metabohzmgussues carneh meme hydranun pruduces pretehs ch2 mo gt hoo3 m Tms creates an arm EHVWDHNEHL Whmh raemtates uhteamhg er uxygen rrerh herhegtemh There 5 Hb uptake at same co2 Hb atse bmds pmtuns thereby pr wmng serhe emeth Must Barbun meme 5 transpunedtu the gms h the farm at mssewee ptasrha bmarbunate ms 5 created W the red menu EHs butmen mfmses nut erthe eu Cu 0 nu M We Lcoqrrm Mo 502 v f g my L c Hurray M H 0 r Fun mm em mm m we quotMI W osoMEwAHn moo Oxygen Uptake and Release Ehmmatton ofCOZ 7 At the guts The hydratmn ruhs backwards as co2 Eaves gms quemy because the ermrehrheht 5 DW h L the rheteeute strhpty ruhs duwngramem There 5 a hse m pH and uptake at uxygen by Hb e The pomt here ts to emphastze the hnkages between 02 and 02 exchange and he r0 e5 of b ood CeHS and p asma Al the Gill 0 cos I WEIKS39 mm Peasma H00 H H J 3 f cho w 90 co Hcor o H39 quotquotquot H CO quot1 002 J Rod H39 Rained H e i 0 Hh quot Mbo2 Hbco Hb o 00 J Bouyancy Gas bladder gas filled sac located between the alimentary canal and the kidneys Filled with coZ 02 and N2 Functions primarily in hydrostatic balance respiration secondarily in sound production and sound reception When used in respiration gas bladder is compartmentalized and highly vascularized Types Physostomous retain connection between the esophagous and gas bladder through a pneumatic duct Physoclistous lose connection Stomach Pnnumalic aim Inles1iiies Types of Gas Bladder esophagus g pneumaiic 39 395 usonimnus uuci Q i piieummlc i 5 mm x stomach y gas y 22352 I rm f 3 539 section pnslull v chamber apeneai m c mm saoium Sucker Seatrout physostomus channel cat sh physoclistous physostomous 91012 Gas Bladder Associated structures in physoclistous shes Antroventral secretory region Gas gland secretes lactic acid lowers blood pH and reduces solubility of dissolved gases A change of1 pH unit releases 50 of O2 bound to hemoglobin raises partial pressure of blood 02 by the Bohr and Root Effects Rete mirabile wondeer net counter current exchange system for gas composed ofa looping bundle ofarterial and venous capillaries Posterodorsal resorptive region Oval thin highly vascularized area circular muscles contract and close the oval preventing gas out ow longitudinal muscles contract and expose the oval permitting gas escape cansmcrm musclu Gas gland Here mlrahlie Wllh parallel vessels mssecrea aoun Gas Bladder As blood leaves the gas gland of the swim bladdervia the venous capillaries of the rete lactic acid is added Reduces hemoglobin39 s affinity for oxygen Oxygen therefore tends to diffuse out and enter adjacent arterial capillaries passing blood to the rete Oxygen concentration builds in the arterial blood as it approaches the gas gland so that the partial pressure of oxygen in the arterial capillaries of the rete is high when it reaches the gas gland Encourages release of oxygen into the gas bladder Arlerlal Muscular sphlncter Here capillary A Oval mirabile venous Lcapillary Swim bladder Gas gland Bouyancy and Blood Chemistry Both physostomes and physoclists have gas glands and associated retia for gas addition How does the gas gland work Requires secretion of gas against a pressure gradient because there is a high concentration and partial pressure of gas inside bladder Blood leading tofrom gland runs through rete mirabile Afferent and efferent flows are in opposition and close together a feature of a countercurrent system Ammal Muscular sphlncrer Flete lt capillary F A Ova mlrabrle venous Leaplllary Swim bladder G35 gland 91012 10 Bouyancy and Blood Chemistry How does the gas gland work 7 There is oxygen in the mood earned by hemogiobtn 39 CeHS in gas giand reiease lactic acid countercunent ow rnarntarns a graorent of hrgh am and tow pH towards gtano The tow pH resutts in oxygen oerng untoaoeo rnto ptasrna by Hi Bohr and Root Effects and a decrease in gas sotuortrty saltingout effect rncreases oartrat pressure of oxygen as t is dtssoived in the ptasrna Counterrcurrent exchange of tons and dtssoived gasses in the rete creates yery hrgh gas pressures in the gas giand facthtattng the ortmsron of gasses rnto the gas gtano Bouyancy and Blood Chemistry 4 m1 1 Least Osmoregulation De nitions 7 Diffusion movement oftons and moiecuies through a rneorurn from a regron of hrgh concentratron to a regron of tow concentration 7 Osmosis rnoyernent ofwater across a semhpermeabie membrane When the concentration ofsoiutes t5 greateron one Side ofthe membrane than the other the net movement ofwaterWtH be from the region of ieSserconcentratton to a regron ofgreater concentratron of rnotecutes 11 91012 Osmoregulation Evolutionarylmplicalions a What about ear y vertebrates m freshwater Osman edmhbnurn Was dtsrupted bythe Law or Osrnosts any amrna submerged m treshmaterthh budy mds somtes m greater cuncentratmn than the surroundth Water tnevttamy tahes m Excess Water Beeorne Water ugged by e Ahsnrplinn through the epdhehum utthe ems and mouth cawty e Smllnwing water atone wtth teed 5d needed a rneehamsrnto getnd orexeess Water Atthe sarne UmE sans are searee m freshwater envtronrnents Eeeause or quSer need a rneehamsrn to retam sans m the budy Sumter vertebrateglomerular kidney Osmoregulation The basic problem for sh 7 TypwcaHy nsh have otrrerent romc concentrauonsman thetr enwonment r they atso have mgmy permeaote membranes 7 Therefore Osmouc movement of worm and Water WM occur 7 Phystotogtcat processes are tmoatreo when tnternat OrHC concentrauons changetartrorn norm Osmoconformers VS osmoregulalors 7 Most nsh are osmoregulators They regmate tnterna mm cuncentratmns to ratny eonstant EVE EXamp E Cahfurma Wh sh Funduus Californianus ean usmuregu ate up to Mme sahmty of seawater and ean surwe up to a1mu514gtlt seawater sahmty 5m an hssuewaml an e 5 plasma nsmmalllv a E SW 79 g g salvan I em 0 m 40 so so 00 um mm m Tmuc um u ml and mum mmrmn r mnmm trumth mum m mtrmmhmn AhcrHldmclhunquLuuncrlIV 39Y01m 39 12 Osmoregulation Osmoconformers vs osmoregulators Most sh are osmoregulators They regulate lnternal lonlc concentratlonsto talrly constant leyel Example Calltornla kllllfl h Hagtlsn ls an osmoconformer lts lnternal lon concentratlon rnlrnlcs the enylronrnent isosmotic 7 Nut surprlslngly tnls nsn lS restncteo to rnanne habltats Osmoregulation in Freshwater Fishes A sh in 39eshwater has body uids in greater concentration than the surrounding water tends to take on water and lose valuable salts to the environment Takes on water by absorbing it through the gills and skin by diffusion Under these conditions the sh exists in a hypotonic environment hypn rneans less39x reters to tne lower solote concentratlon ottne surruundlng Water 7 Problem is to eliminate water and retain salts Drlnklng yery llttlE water Has numeruus large Wellrdeyeluped glumerull Reabsurbs salts alung tne length or lts conyoloteo toboles Pruduces large amuunts or a yery ollote onne 542 or body Welght oay Osmoregulation in Freshwater Fishes absorb water through align 5km i l abtalns sails inrougn chluvlda39 calls to gllls and mm loan lest wa leans lemmas moan water and some salt yla dllule mmn Open arrows lnolcate rnoyernent or substances by passlye dl usl n closeo arrows lnolcate rnoyernent of substances by actlye transport rnecnanlsrns 91012 13 91012 A closer look at the kidney F suwngvnmv Functional components Glomerulus a typical WM kidney of a freshwater sh has tens ofthousands of large glomeruli Large m mnyw my amounts ofwater pass 7 u NMWMWW through them Provides a M ltrate that can be modi ed selectively by the kidney tubule 039 am at Neck Region lined with cilia ciliary action aids movement of materials into tubule Important in the lowpressure ltration systems of shes First Proximal Segment PCT I location of reabsorption of many macromolecules eg hunvvzt39uiv glucose proteins also 37quotm quot75339k excretion of organic acids mmw J smmm mssanmmmsv A closer look at the kidney J L FanialoanunmnsLu Functional components lama Second Proximal limbo T l mac Segment PCT ll largest my Wth any region of tubule has high ka metabolic activity ie active transport N Wm w Wm mechanisms that are l l Q df c responsible for reabsorption r mg mm ofmany saltsegMg1 Per N m sorCa139 PNa Cl39 and quot3 A Hcoa u gt23 so 6 p Intermediate Segment highly ciliated portion that assists in moving uids imam through the tubule ln D Wgw 391 quot3 5 freshwater sh it is w or important to move the tluid cu c I through the length ofthe tubule as fast as possible to 2mm minimize reabsorption of WM quot 9 water OAmeltm lemuw rassnwnsn mam FawnSUV mman My A closer look at the kidney l Functional components will wquot Distal Segment ncn 39 quot participates in active reabsorption of Na and 7 imam wastns some Cl39 My a Longitudinal Collecting l Glucose mums Duct CT reabsorbs ncr N Cl m0 mostly Na and some Cl39 monovalent ions again L ll lntemmmail 5mm my We luv 7 m wl m m Z mnw tls k9 w m m5 Actw llow omwuw 14 Osmoregulation in Freshwater Fishes A closer look at the kidney 7 Functlonai components 7 End result Dllute urlne contalnlng mostly WatEr but also some aikaiulds some amlno acres and a llttle urea and ammonla Altnougn some nltrugenuuswaste ls excreted ln tne urlne amuunts to uniy 745 ortotal nltrugen excreted byfresnwater sh e Must passesuui inmughihe gliisasammunla Osmoregulation in Freshwater Fishes Retaining salts e Kldney alone cannot reabsorb enougn salts to malntaln osmoreguiarlty e Chloride cells 7 speclal cells ln tne ngS and oral membranes to absorb long by actlve transport mecnanlsms Absorbs acld pnuspnate brurnlne caiclurn cnlonde ilthlum Sudlum suifate iDrlS etc Osmoregulation and Excretion mm s 91012 15 Osmoregulation in Marine Fishes Marine shes have a total quantity ofsalts that is less than that of the same volume of seawater Some marine teleosts have as little as onethird the osmotic concentration of seawater Because their body uids are less concentrated than seawater they tend to lose waterthrough their membranes Under tnese condltlons tnensn exlsts ln a hypertonic environment 7 quotnvperquot means quotmore39x rerers to nlgner solute content ln surruunulng water Osmoregulation in Marine Fishes Marine shes have a total quantity ofsalts that is less than that of the same volume of seawater Some marine teleosts have as little as onethird the osmotic concentration of seawater Because their body uids are less concentrated than seawater they tend to lose waterthrough their membranes Problem is to conserve water and excrete excess salts 39 Drlrlk seawater 39 Have fewer and smaller glomerull Excrete salts alongtne lengtn ottne convoluted tubules Produce small amounts of averv concentrated unne as llttle as 2 5 ml per kg of body Welghtday Osmoregulation in Marine Fishes mediumta rum momkg losnswnler may llnmsce doc mOswl kg lmougn orls r I f w m 5km l l w A t lm removes sails vla clvlnnoe cells ml gulls aims water and salts tsv swallowmg Eeawnlcr and teen sails lost y vta laces salts and llllle waist lost on scam um Open arruws lndlcate movement or supstances pv passlve ulqulerl closed arruws lndlcate movement or supstances pv actlve transport mecnanlsms 91012 16 A closerlook attne kidney 7 Functionalcomponents Glomerulus glomeruliin marine teleosts are SmaHi poorly yascularized and blood pressure is low May be lost aglomemlar Neck Region may be lost altogether especially in aglomerularspecies First Proximal Segment PCTI location of reabsorption ofmany macromoleculese 9 glucose proteins also excretion oiorganic acids A closerlook at tne kidney 7 Functionalcomponents Second Proximal Segment PCTII instead ofactlve reabsorption of salts as in freshwater teleostsi tnis is the site of actiye secretion of salts e 9 ngla soy Cain Pi NanClxandHcoa Also responsible for actiye secrection oinitrogenous Wastes urea creatine creatinine lntennediate Segment absent in marine flSH because tne need nere is to slowtne movementof uld to maximizethe amount waterpassiyely diffusing backll ltothe blood A closerlook attne kidney 7 Functionalcomponents Distal Segment DS participates in some reabsorption ofNa and Cl Longitudinal Collecting Duct CT some reabsorption ofNa and some or Mminzmausr a mi 72 team mam oi ii i in W m iiimywmy Neat ulnamwaeite i cum m iio chl H to poi in so so i o inimaaai 7 mama meosr new J53 imam w era ca is now in may iiiNew waSV 3 an amine n mi amid c wmm lls am 7 ii i lnlmnme uir Halt wt may ui iw W 300 mm ui 39 unm viii Mllvu Hm iqm v cw MARINE iamsr C7 naea 350nm Me imamiii is Lenin Na a me We a I an m NaiCl imam 91012 17 Osmoregulation in Marine Fishes A closer look at the kidney Functional components End result A small volume ofstrong concentrated urine is produced containing creatlnlne creatlne some urea some ammonia plus other nitrogenous compounds Again noweven 90 of nitrogenous Waste is eliminated by tne gills as ammonia and urea Excreting salts Kidney alone cannot eliminate all excess salts to maintain osmoregularity Chloride cells special cells in the gills and oral membranes to secrete ions by active transport mechanisms Osmoregulation in Elasmobranch Fishes Marine elasmobranchs osmoregulate in a very different way 39om teleosts Evolved a specialized segment of the nephron that reabsorbs urea and returns it to the blood In ux of urea and TMAO trimethylamine oxide raises the osmotic pressure of the blood to a level just above that of seawater so that water ows into the body of the shark similar to freshwater sh Have numerous welldeveloped glomeruli and excrete large amounts of dilute urine Osmoregulation in Elasmobranch Fishes medium ca 1000 mOsmrkg body tlulds cs nun mDsmkg gills mack lass a ma am TMAO mas urea and mm mlained by kidney divalenl Ions excrelad n urine lngesis sans wun icon water absorbed by gills and skin Sodium exclalea quot5 by recial gland ins vxa laces 91012 18 91012 Thermoregulation Ectothermy Advantage low metabolic costs requires less energy and less food Disadvantage cannot live or function very well in extreme thermal environments especially the cold 0 Endothermy Advantage biochemical reactions become more efficient fish can utilize wider thermal ranges Disadvantage high metabolic costs requires more food leaves less energy for reproduction growth etc Thermoregulation 0 Heat Conservation Regional Endothermy different temperatures in different parts of an animal39 s body Rete mirable near large swimming muscles often found in tunas and some sharks Fine counter current network of veins and capillaries that Rm mimbiln M mm Culmm yum Emma s new my cam Thermoregulation 0 Heat Conservation Regional Endothermy Rete mirable near large swimming muscles Rete mirable on the liver Functions in maintaining increased gut temperatures aids in digestive ef ciency also found in tunas and sharks Warm parts of the central nervous system especially brain and eyes Allows fish to use deeper colder more biological productive habitats without a decrease in brain and visual function Modify the circulatory system by having retes near the eyes and brain better vision at cold temperatures 20 Enmnigyg ggvgenei Psaudabrznchla Thermoregulation Jljr Lenomuameie wearer V3 Shiba k 39 MJ 921 asym fkg 34 raidz qg m Wares lmmnm cavam Plexus my mm Psmaomancmal m Flight omual Hale mme Amanor Tubbesing V A and B A Block 2000 Orb tal rete and red muscle vein anatomy indicate a high degree of endothermy in the brain and eye of the salmon shark Acta Zoologica 81 49 56 Thermoregulation Heat Conservation Regional Endothermy Thermogenic tissues special heatproducing tissues Billfish have specialized eye muscles without contractile filaments that produce heat without muscle contractions when stimulated by nerves FIGURE 71 Stimulation olrhe modi ed musde ell ofrhe bill m mam holidnal melabairsm generating adriftanal heal iewu imulusE meiaboiisr l MI ocuorm m Hualul Cull 91012 21
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