98

CHNG VIII. TRAO I CHT V DINH DNG

A. 1. Trao i cht ngha sinh vt hc ca trao i cht

Trao i cht l c trng c bn ca c th sng. l im c bn phn bit gia sinh vt v phi sinh vt. Trao i cht ngng th s sng khng cn na. Trao i cht l s s dng sinh hc cc cht dinh dng hp thu c cho qu trnh tng hp (chng hn cho sinh trng) v tiu ph nng lng. Trao i cht bao gm hai qu trnh c bn tri ngc nhau v c lin h mt thit vi nhau l ng ha v d ha. ng ha l qu trnh bin i nhng vt cht hu c c cu to n gin thnh nhng vt cht c cu to phc tp xy dng c th v to ra ngun nng lng d tr. D ha l qu trnh bin i nhng vt cht cu to c th v ngun nng lng d tr c cu to phc tp thnh nh ng vt cht n gin v phng thch nng lng m c th c th li dng c. ng ha v d ha hon thnh c l nh hai phn ng phn gii v tng hp. Nhng vt cht dinh dng u c tim tng nng lng nht nh nn trong qu trnh vt cht bin i th km theo s bin i v nng lng v th trao i cht bao gm hai mt khng tch ri nhau l trao i vt cht v trao i nng lng. Trao i cht li bao gm hai ni dung: (1) s trao i vt cht v trao i nng lng gia c th vi mi trng; (2) S chuyn ha vt cht v nng lng bn trong c th. 2. 2.1 Trao i cht Trao i cht protein

Protein l thnh phn quan trng nht ca mi c th sng. N chim khong 16 18% trng lng c th ng vt v trong c th n trng thi cn bng ng . Cn bng ng c ngha l cc sn phm trung gian ca qu trnh trao i cht protein tuy c th tm thi n nh nhng v cht th khng ngng i mi. Khc vi cc hp cht hu c khc cng tham gia to thnh c th, trong protid ngoi carbon, hydro, oxygen ra cn c nitrogen, sulfur v i khi c c phospho. Protid ph bin nht l albumin v globulin cha khong 54% C, 7% H, 16% N, 1% S v 0,22% O . Mi loi ng vt c cc protein chuyn bit ca n . 2.1.1 Chuyn ha protein trong c th

Protein trong thc n sau khi tiu ha c hp thu vo mu di dng cc acid amin ri qua tun hon tnh mch ca vo gan. Sau khi c hp thu vo c th, cc acid amin thc n cng vi cc acid amin t qu trnh d ha protein trong c th to thnh tp hp trao i cht cc acid amin. Cc acid amin c chuyn ha theo cc hng sau:

SLVTS

NVT

99

- c tng hp thnh protein mi trong c th, bao gm c protein huyt tng v hemoglobin. cc ng vt trng thnh cc protein lun lun c thay th vi nhng t l khc nhau. chut 1/2 tng s protein c thay th trong 17 ngy, ngi l 86 ngy. - To thnh nh ng cht c bit c bn cht protein nh cc hormone, cc nucleic acid, cc enzyme. - Nhng acid amin dng lm ngun nng lng s c kh amin to thnh keto acid, t keto acid s to thnh glycogen nh ngun d tr nng lng tm thi v thnh lipid nh ngun d tr nng lng lu di. Keto acid cng c th xi-ha thnh CO2 v H2O gii phng nng lng cho c th s dng. Ngoi ra phn c m (gc amin) c th bin i thnh ammonia, urea hoc uric acid thi ra ngoi c th. - Thng qua tc dng thay i gc amin, gc amin c th chuyn sang cc hp cht khc to thnh acid amin mi. - Qua tc dng kh gc COOH to thnh amin, b ng con ng ny tyrosine chuyn thnh tyramin v histidine thnh histamin, l nhng cht c hot tnh sinh hc. - Mt phn acid amin c th thi theo nc tiu v phn. 2.1.2 Cn bng nit S cn bng nit c biu th b ng cng thc sau: N n + n =k

N: lng nit ly vo qua thc n n: lng nit thc n khng hp thu c n: lng nit b phn gii v bi tit ra ngoi - Khi k = 1 N = n + n c gi l trng thi cn bng u xy ra nhng c th trng thnh. - Khi k > 1 N > n + n c gi l trng thi cn bng dng xy ra trong thi k c th sinh trng, pht dc, hi phc sc khe sau khi m hay sau khi i. - Khi k < 1 N < n + n c gi l trng thi cn bng m xy ra trong thi k c th gi ci, au m, thiu dinh dng hay l b i lu ngy. 2.1.3 Vai tr ca gan trong trao i cht protein - Tng hp protein cho c th. Cc lo i protein huyt tng (prothrombin, fibrinogen) ch yu c tng hp trong t bo gan.

SLVTS

NVT

100

- Gii c cc sn phm phn gii protein. Gan c tc dng bin i NH3 (ammonia) do qu trnh kh amin thnh urea c loi thi trong nc tiu. Nu c nng ca gan b ph v, urea khng hnh thnh c, lng ammonia tng ln ng vt s b ng c m cht. 2.1.4 Vai tr ca protein trong c th

Khc vi lipid v carbohydrate, acid amin trong c th khng c kh nng d tr v cng khng c ch d tr nht nh. Acid amin tha trong tp hp cc acid amin b kh amin v sn carbon c xi-ha to nng lng hay bin i thnh carbohydrate hay lipid. - Protein c vai tr v cng quan trng i vi c th, n l thnh phn cu to ca c th (l thnh phn ch yu to nn nguyn sinh cht t bo), l nguyn liu b sung v pht trin c th. C th ni qu trnh sinh trng ca c th l qu trnh tng hp protein mi trong c th. Mt s loi acid amin nh tryptophane, lysine, histidine... rt cn thit cho s sinh trng c th ng vt m b n thn chng khng t tng hp c, phi da vo s cung cp t thc n. Nu thc n thiu cc thnh phn acid amin th sinh trng ca ng vt b st km. Nhng cht ny gi l acid amin thit yu hay cn gi l acid amin khng thay th c. Protein no cha y cc loi acid amin gi l protein c gi tr ho n ton. Nhng protein no cha nhiu loi acid amin thit yu th c gi tr dinh dng cao hn. - Protein cn l ngun nng lng. Mi gram protein c xi-ha trong c th s sinh ra mt nng lng l 4,25 kcalo. 2.2 Trao i cht lipid

Trong ng tiu ha m ca thc n c phn gii thnh glycerol v acid bo. Khi vo t b o biu m ca mng nhy ru t chng li hp thnh m trung tnh ri i vo h bch huyt, mt phn nh i vo mu (30%). 2.2.1 S chuyn ha lipid trong c th Khi vo c th m c chuyn ha theo cc hng sau: - D tr li di dng m d tr, ch yu di da v cc m lin kt ni tng. Cc loi c khc nhau th ni tch ly m cng khc nhau. C sn m tch ly gan nhiu nht, c chp tch ly m mng ru t, c chy tch ly m nhiu nht t chc lin kt di da, c lo i tch ly m c. - Tng hp nhng thnh phn cu to t bo ca cc t chc c th (phospho lipid).

SLVTS

NVT

101

- Phn gii thnh glycerol v acid bo sau trc tip oxi ha thnh CO2 v H2O ho c chuyn bin thnh glycogen gan. - c cc tuyn th s dng to nn thnh phn ca cht ni tit c bit nh cc hormone steroid. 2.2.2 Vai tr ca gan trong chuyn ha lipid M trong gan rt hot ng v mt trao i cht. Ba tc dng ch yu ca gan l: - Lm cho acid bo bo ha thnh khng bo ha to iu kin thun li cho s phn gii tip tc hoc chuyn ha ca chng. - To thnh cc vt cht phospholipid tham gia xy dng cc m (mng t b o, nguyn sinh cht) hay b xi-ha. - To thnh cc th ketone l sn phm trung gian ca chuyn ha m. Sau khi c to thnh gan cc th ketone ny s c chuyn n cc m, nht l m c, xi-ha v sinh nng lng. 2.2.3 Vai tr ca lipid trong c th

- Lipid l nguyn liu c nng lng cao nht, 1 gr lipid khi c xi-ha s sinh ra 9,45 kcalo. Lipid li c th d tr nhiu trong c th nn c ngha quan trng v mt d tr nng lng . - M l dung mi ca nhiu vitamin (A, D, E, K). Khi n nhng thc n c m cng thng c cc lo i vitamin ny. - Cc acid bo c th c to thnh trong c th do mt lot phn ng phn gii v tng hp acid bo, chiu hng ca qu trnh ny ph thuc vo nhu cu nng lng ca c th. Tuy nhin, cc acid linoleic, linolenoic, arachidonic v mt s acid khc c l khng th tng hp c trong c th ng vt, l nhng acid bo khng th thay th ch c trong thc vt. Khi thc n thiu nhng acid ny th s trao i m, thnh thc sinh dc, ... b ri lon. - M l mt trong cc thnh phn ca t chc, c bit phospholipid l thnh phn quan trng ca mng t b o v nguyn sinh cht c lin quan n tnh thm thu ca t bo. 2.3 Trao i cht carbohydrate (COH)

Cc ng a (polysaccharide) trong thc n sau khi tiu ha bin thnh cc ng n (monosaccharide). Khi monosaccharide c hp thu vo mu th bin thnh ci gi l ng mu. Qua s tun hon ca mu , ng mu phn b n khp b phn trong c th. 2.3.1 S chuyn ha ca COH trong c th C 3 bc chuyn ha:

SLVTS

NVT

102

- Mt phn c cc m trc tip s dng lm nguyn liu cung cp nng lng. - Mt phn c tng hp thnh nng lng d tr tm thi glycogen , ch yu gan v c. T l hm lng glycogen ca gan v c l 1:7. - Mt phn ln glucose chuyn ha thnh lipid nh ngun nng lng lu di. 2.3.2 Vai tr ca gan trong chuyn ha COH

- V tt c ng vt s dng glucose nh mt ngun nng lng cho cc hot ng ca bn thn chng nn gan c mt vai tr quan trng trong s trao i cht ng. Trc ht, n duy tr mt ngun quan trng ca glycogen vt qu nhu cu trao i cht ca ring bn thn n, sn sng chuyn ha thnh glucose t do bi qu trnh glucogenolysis tc l s phn gii glycogen duy tr hm lng ng mu. V phng din ny glycogen gan tc ng nh mt ngun carbohydrate cho to n b c th. Sau khi glucose thc n i t rut vo mu v nng glucose mu gia tng, gan h p thu mt s glucose ny t mu v bin i n thnh glycogen, v cng vi cc c quan khc nh c, no v cc m khc lm cho hm lng ng mu dn dn gim xung. Khi glucose mu gim xu ng di mt mc no , mt phn ca glycogen gan c phn gii v gii phng vo trong mu l glucose t do v bng cch ny hm lng ng mu c gi trong nhng bin ng nh. - Vai tr quan trng khc ca gan trong trao i cht ng l tng hp glucose mi (gluconeogenesis). Trong khi i cc c c th s dng m d tr nh mt ngun nng lng nhng h thng thn kinh th ph thuc vo glucose d n xut t acid amin c to ra bi s phn gii cc protein m.
Protein t.a

~3,8 Kcal/g
Chc nang D tr

Tieu hoa

COH t.a

1,2-3,9 Kcal/g
MAU & GAN

Tieu hoa

Tie u hoa

Lipid t.a

~8,5 Kcal/g
Cau truc

A. acids + peptids Amino acids 2-3 C.

Protein

Cac hat m Ca c acid beo Cac monoglyceride

Glucose 6 C. (Can insulin)

D tr Lipids

2,5-4% BW NH3 (Bai tie t)

Kh amin

Pyruvate 3 C. Lactate + 2 ATP

1-30% BW O2
G a n

CO2

Glycogen (phan t 5 glucose) 30 C.

C Carbohydrates

Oxi hoa

Ghi chu

3 C. = chuoi 3 carbon % BW = % Tro ng lng t T le glycogen gan:c = 1:7

Acetate 2 C. O2 ho hap
Chu trnh citric acid

2-4% BW

CO2 + H2 O + 33 ATP

H.41 Cc con ng trao i cht chnh ca cc protein, carbohydrate v lipid c

SLVTS

NVT

103

2.3.3

Vai tr COH trong c th

COH l ngun nng lng ch yu cho mi hot ng sng ca c th. Mi gr glucose khi xi-ha hon ton sn sinh ra 4,25 Kcal. Khi thc n cha COH tng ln th s phn gii lipid v protid trong c th gim i, v nng lng ch yu do glucose cung cp. Glucose sau khi c hp thu vo mu s chuyn ha thnh glycogen nh ngun d tr nng lng tm thi v phn ln thnh lipid nh ngun d tr nng lng lu di. COH l mt trong nhng thnh phn khng th thiu c trong cu to ca t chc c th nh cc cht glucoprotein mng t bo. 2.4 Trao i cht nc

Nc trong c th c t l cao nht, ng vt cao ng c khong 7075% nc so vi trng lng, lo i c c khong 8085%. Ni chung t l tng i n nh, song s n nh ny cng l mt loi cn bng ng, ngha l nc trong c th lun lun mt i nhng li c b sung khng ngng. Nc trong c th mt phn nh dng t do, nhng phn ln dng lin kt; v d lin kt vi protid thnh th keo. Trng thi lin kt ny lm cho nc mt tnh cht chuyn ng v sc ha tan. Trong tnh trng mt nc s to nn s rt nc t chc ng thi cng s dn n mt nhiu mui nh hng n s cn bng acid-base, dn n tnh trng ng c acid. Nc trong c th c ngha quan trng v sinh l. Bi v: - Nc l thnh phn quan trng ca ni mi trng. Nu nc gim xung th nng thm thu ca mu tng ln, mu chy chm chp khng th cung cp kp thi cc cht dinh dng cho cc c quan, tr ngi n s trao i cht ca cc t chc. - Mi phn ng sinh ha trong c th u xy ra trong nc. Thiu nc s trc tip nh hng n qu trnh sinh ha ca c th. - T nhit ca nc tng i cao v nc c th d tr c nhiu nhit nn c th ngn chn s bin i t ngt ca thn nhit, duy tr s hot ng b nh thng v trao i cht trong c th. Vai tr quan trng ca nc c th c minh ha, v d khi c b i lu, m d tr hon ton b tiu bin nh ng c vn sng; song ch cn mt 10% nc ca ton c th, c s cht. C l ng vt thy sinh, dch c th c p sut thm thu rt khc vi mi trng bn ngoi nn thng xuyn c s trao i nc gia c th vi mi trng. Thng qua ho t ng ca thn, c c th iu ha qu trnh trao i ny. 2.5 S trao i mui khong

Ngoi cc cht ch yu nh m, ng, m c to thnh t carbon, hydro, oxygen v nitrogen ra, trong c th cn c kh nhiu nguyn t to thnh mui v c cung

SLVTS

NVT

104

cp cho nhu cu ca c th. Vai tr ca chng khc nhau, c loi l nguyn liu to n n t chc, c loi l cht cn thit duy tr ch c nng sinh l bnh thng, c loi l cht xc tc phn ng ha hc. - l nguyn liu ca t chc nh cc mui phosphate potassium v carbonat potassium l thnh phn quan trng ca xng v rng. St l thnh phn quan trng ca myoglobin v hemoglobin; phospho l thnh phn quan trng ca phospholipid... Tt c u l nhng thnh phn khng th thiu c duy tr s hot ng b nh thng ca c th. - loi duy tr chc nng sinh l b nh thng nh cc loi mui kim gip cn bng acid-base, duy tr p sut thm thu v cn bng nc, v.v. - l cht xc tc nh trong phn ng trao i cht COH, Mg2+ c th thc y cho phn ng tin hnh thun li. Mt s cc kim loi l cht hot ha cc enzyme tiu ha... - cc mui khong c vai tr c bit quan trng trong s dn truyn thn kinh v l thnh phn cu to ca cc hormone c tc dng iu ha qu trnh trao i cht b nh thng ca c th di nh hng ca cc yu t bn trong cng nh bn ngoi. c thng xuyn c s trao i mui khong vi mi trng b n ngoi. C thng qua ho t ng ca mang, rut v thn iu ha cc qu trnh trao i ny. + Tc dng sinh l ca m t s nguyn t vi lng Nguyn t vi lng tn ti trong c th vi mt lng rt nh (10 -310 -12%) nhng c nh hng r rt n qu trnh trao i cht ca c th. Nhng nguyn t quan trng nht l st (Fe), ng (Cu), cobal (Co), iod (I), mangan (Mn), km (Zn) v fluor (F). - St (Fe): Fe trong c th di dng hp cht hu c (hemoglobin, myoglobin...) v hp cht v c (Fe d tr). St d tr ch yu trong gan sau l t tng v ty xng. St trong c th nhiu dng khc nhau v phng din kt hp ha hc v v phng din chc nng. St l mt thnh phn ca hemoglobin gi vai tr vn chuyn oxygen. - ng (Cu): Cu l b phn to nn nhiu enzyme xi-ha nh polyphenol oxidase, lactase, acid ascorbic oxidase v tyronase nn n c lin h cht ch vi h hp m bo. Cu l cht xc tc to thnh Hb, thc y s s dng st. Thiu ng th s bin dng Fe cng b nh hng, ng vt xut hin triu chng thiu mu c tnh cht dinh dng, sinh trng ngng. ng tham gia to thnh sc t en. Lng Cu cn thit vo khong 6/7 Fe. - Cobal (Co): Co l mt b phn to thnh vitamin B12. c chp, s b sung Co trong thc n lm gia tng s lng hng cu v hm lng Hb, gim t l cht, gia tng sinh trng v sinh sn v gia tng s li d ng thc n ca c.

SLVTS

NVT

105

- Iod (I): hm lng iod trong c th rt t, a s cha trong tuyn gip, I tham gia to thnh Iodotyrosine v thyroxine c phn b rng ri v ch c ng vt xng sng. Iod thng qua hot ng ca tuyn gip kch thch trao i cht ca c th. Nu thiu I ng vt b cc ri lon v sinh trng v thnh thc sinh dc. - Mangan (Mn): Mn l cht kch thch ca nhiu enzyme trong c th, n c nh hng trc tip n vic tch ly Ca, P v thc y tc dng to xng. Khi trong thc n hng ngy ca ng vt non khng Mn th hm lng enzyme phosphatase trong mu v xng gim nh hng n ha ct xng ca con vt, xng b bin hnh. ng vt trng thnh thiu Mn th chc nng sinh dc s suy yu. - Km (Zn): Zn l thnh phn cn thit ca enzyme carbonic anhydrase, cht xc tc s hydrat ha ca CO2 trong nhiu m nh mang, t bo mu v thn ng vt xng sng cho nn Zn l nhn t cn thit cho qu trnh h hp ca m bo v cn bng acid-base ca thn. - Fluor (F): F c tm thy trong cc cht n v xng v rng nhng trong s thng d gy ra nhng cu trc khng b nh thng. F c th c ch mt s enzyme do khi F qu nhiu s nh hng n trao i cht. 2.6 Vitamin v s trao i cht

Vitamin l nhng cht hu c cn thit cho c th. Chng khng phi l nhng nguyn liu ch yu to nn c th, cng khng th cung cp nng lng nhng li l mt trong nhng thnh phn ca nhiu enzyme quan trng tham gia vo qu trnh trao i cht. Lng vitamin dc i hi rt t nhng v phn ln ng vt khng t tng hp c nn phi ly t thc n. Khi c th thiu hay tha vitamin th ng vt s mc nhng lon chng c bit cc bnh thiu hay tha vitamin. 2.6.1 Vitamin ha tan trong nc

Cc vitamin ha tan trong nc c chc nng nh cc coE trong cc phn ng trao i cht chuyn bit, cn thit i vi hu ht, nu khng ni l tt c, t bo ng vt. 2.6.2 Vitamin tan trong m

- Cc vitamin A: c bit di 2 hnh thc A1 v A2. Vitamin A1 hin din ng vt xng sng cao ng v c bin, cn A2 chim u th c nc ngt. Hai phn t vitamin A c to thnh t mt phn t carotene, c trong nhiu thc vt, xy ra gan, nhng chnh rut. Trong s thiu ht trm trng n c th gy ra s tr hon s pht trin xng v lm tn thng da c. D tha vitamin A trong khu phn c th gy c, triu chng l sinh trng gim, microhematocrit b h thp v s b o mn vi ui v cung ui. - Vitamin D: c nhiu hnh thc D1, D2, D3

SLVTS

NVT

106

D3 hin din gan c gip s hp thu Ca t ng tiu ha, v vi parahormone iu ha mc canxi mu v s hot ha Ca cho s khong ha xng. - Vitamin E: s thiu ht vitamin E gy ra mt s thoi ha ca biu b tinh nang cho ra nhng tinh trng khng hot ng. S hoi t gan, xut huyt cc b v tn thng tinh so c quan st c. - Vitamin K: hin din vi 2 hnh thc K1 v K2 tht c c K2, kch thch s to thnh prothrombin cn thit cho s ng mu. S thiu ht dn n s xu t huyt. B. 1. Nng lng sinh hc (bioenergetics) Tng quan

Bioenergetics l nghin cu vic s dng nng lng bi cc sinh vt sng. i vi ng vt, nghin cu ny bao gm phn tch cc ngun nng lng, cc phng php thu nng lng, cc con ng phn b ca n trong con vt, cc cng s dng di nhng iu kin khc nhau v trng thi nng lng cui cng ca cc sn phm ang ri khi con vt. Cc n v ca nng lng thng l calorie (cal) hay kilocalorie (Cal hay kcal) v joule (J) hay kilojoule (kJ) trn mt n v trng lng chuyn bit. Cng tiu hao (s dng) nng lng thng c tnh kcal/kg/gi hay kcal/kg/ngy. Vic o nng lng tiu hao trc tip vi sn xut nhit th kh p dng vi cc ng vt mu lnh nh, chng hn nh c, nn nng lng tiu hao lun lun c o gin tip thng qua tiu hao oxygen. Tiu hao oxygen i khi c tnh bng mL O2/kg/gi (vi th tch oxygen c iu chnh theo nhit v p lc khng kh) nhng thun tin nht l tnh b ng m g O2/kg/gi. Gia th tch v trng lng oxygen c th chuyn i: 1 mg O2 = 0,70 mL O2; v gia oxygen v nng lng: 1 mg O2/kg/gi = 0,00337 kcal/kg/gi hay 0,081 kcal/kg/ngy, 1 kcal/kg/gi = 297 mg O2/kg/gi; v 1 kcal = 4,184 kJ. i vi cc ng vt, ngun nng lng c bn l thc n nhng nng lng thc n s khng c gi tr cho ti khi thc n c n vo, tiu ha v ng ha bi h thng tiu ha. Phn ln nng lng c gii phng t thc n bi qu trnh xi-ha nn nng lng sinh hc thng c kho st di dng tiu hao oxygen. Cc kha cnh phn t ca dng nng lng (energy flow), mt ni dung ca trao i cht (metabolism), l mt s kt hp chc nng ca gan v c, nhng cng lin quan n s iu ha thm thu v bi tit cc sn phm tha ca b my trao i cht. Mt s lng nng lng v vt cht th khng cn cho cho duy tr v cc yu cu thng thng cho ho t ng sng thng nht s c s dng cho tng trng, sn xut cc sn phm sinh d c v d tr. Alexander (1967) din t tng chung ca s phn b nng lng v s quan trng ca vic s dng nng lng hiu qu trong mt cng thc: F = g (G + H) + R + S

SLVTS

NVT

107

Trong : F: lng thc n n vo G: sinh trng (sn xut m mi) H: cc giao t R: trao i cht c bn S: bi hay hot ng khc : hng s, thng bng 0,8 (20% thc n b mt trong phn, nc tiu, hoc ammonia thi bi mang m khng i vo cc con ng sn xut nng lng trong qu trnh trao i cht ca c) Yu t tng trng v sn xut giao t, g, xp x bng 2 bi v cn gp i nng lng sn xut cc m mi so vi vic duy tr cc m hin ti. Mt s t l phn trm tiu biu cho vic s dng thc n trong cng thc ca Alexander nh sau: 0,8 x 100 = 2 (5 + 1) + 34 +34 C nhng thi im lm cho cc gi tr trn khng chnh xc, chng hn c khng n khi di lu hay sinh sn. Mt th d khc l c bn (flounders) c cho n mt khu phn n ti thiu vn tip tc sinh trng nhng khng sn sinh giao t, H = 0. Hay c hai cng vn ng (S) v trao i cht c bn (R) c th thay i theo nhit v ma v. ngha ca cng thc trn l ch ra nh hng tng i ln trn s sinh sn (H) ca nhng thay i tng i nh trong bt k thay i ca cc yu t khc. V d, nu thc n n vo tng t 100 ln 101, s sn xut giao t tng t 1 ln 1,4 (40%). Hay nu c n lng thc n bnh thng nhng gim vn ng 2% (S = 32) th H s tng ln 2 (100%). Sinh trng v sinh sn dng nh l tng s v l s kt hp ca mt lch s qu kh ca c phi hp vi kh nng i ph vi mi trng ca n. 2. Bao hot ng (performance envelope) ca c

Mt cch din t nhng kh nng nng lng sinh hc ca mt con c l vi mt a gic 4 cnh (bao ho t ng). Cc cnh phi v tri l nhng gii hn nhit gy cht cao v thp cho mt con c no trong khi cc cnh trn v di l cc cng tiu hao oxygen ti thiu (ng vi trao i cht c s hay tiu chu n) v ti a (hot ng tch cc n nh). Bin gii trn thng gm 2 phn, phn bn tri c cho l b gii hn bi nhit trong khi phn bn phi dng nh b gii hn bi

H.42 Cng tiu hao oxygen ca 4 loi c trong s lin h vi nhit (theo Brett, 1972)

SLVTS

NVT

108

tnh gi tr ca oxygen. Phn ln, nhng khng phi l tt c, c cho thy c s gin on tng t v cng tiu hao oxygen cc i. Phn ln cc cng tiu hao oxygen trong mt ngy tiu biu trong i sng ca mt con c xy ra trong cc bin gii bao hot ng ca n v phn ln l na di ca gii hn c bn v tch cc. Gii hn hot ng ca c th tng t nh lng c v b st nhng thp hn chim v hu nh. Phm vi trao i cht (metabolic scope) l t s gia cng tiu hao oxygen tch cc v c s. Cc loi c khc nhau c phm vi trao i cht khc nhau. 3. Trao i nng lng

Vt cht dinh dng u c cha nng lng nn qu trnh trao i vt cht cng km theo qu trnh trao i nng lng. Cc qu trnh trao i nng lng rt phc tp, hnh thc a dng nhng cui cng u bin thnh nhit nng thi ra ngoi. 3.1 Gi tr nhit ca cc cht dinh dng

Mi gram COH hoc lipid d t chy bn ngoi hay bn trong c th u sinh ra mt nng lng Q = 4,25 Kcal v 9,45 Kcal. Mi gram protein khi t chy bn trong c th sinh ra mt nng lng Q = 4,25 Kcal (khi t chy bn ngoi c th ta ra 5,4 Kcal). Tr s nng lng ca cc cht dinh dng khi t chy bn trong c th c gi l gi tr nhit ca thc n. Mc d nhng gi tr ny c dng cho tt c cc loi ng vt trong nhiu nm (v vn cn trong nhiu nghin cu) gn y chng c iu chnh ty theo kh nng tiu ha ca con vt c nghin cu. Phillips and Brockway (1959) xc nh cc gi tr nhit i vi 3 lo i c hi (brook, brown v rainbow trout) nhn thy chng c kh nng tiu ha v hp thu 85% i vi lipid, 40% i vi carbohydrate v 90% i vi protein. S dng cc mc tiu ha ni trn v c s iu chnh cho protein i vi hm lng nit ca n (nitrogen khng th c gi tr v nng lng), cc tc gi xc nh gi tr nhit i vi c hi l 3,9 kcal/g protein, 8 kcal/g lipid v 1,6 kcal/g COH. Cc gi tr ny khng c nh i vi cc lo i c khc tr phi chng tiu ho cc nhm thc n c nh gi tng t nh c hi. 3.2 Tnh ton nng lng thc n

Nng lng thc n c xc nh bi nhit sinh ra trong s t chy hon ton trong mt nhit lng k (calorimeter). n v tnh l calorie (cal) hay kilocalorie (Cal, kcal) v joule (J) hay kilojoule (kJ) l s lng nhit cn thit nng nhit 1 kg nc ln 1 oC. (1 kcal = 4,184 kJ). C hai phng php c dng tnh ton gi tr nng lng ca khu phn thc n.

SLVTS

NVT

109

(1) Nng lng c xc nh mt cch trc tip trong mt bm nhit lng k (Maynard v Loosh, 1962). (2) Nng lng tng cng ca khu phn n c xc nh gin tip t gi tr nhit trung bnh ca cc nhm cht dinh dng (protein, lipid, COH). Phng php th hai thng c dng v phng php th nht th lu, phc tp v tn km. 4. 4.1 Cc yu t ch yu nh hng n trao i cht c th Trao i cht c s (tiu chun)

Vi mc ch thc tin nh gi hay so snh cng trao i cht ngi ta thng dng khi nim trao i cht c s. Trao i cht c s l s trao i cht ca ng vt trong iu kin tiu chun. ng vt tnh trng yn tnh, d dy khng c thc n, thn kinh khng b cng thng v nhit mi trng ti thch hp. Nng lng do trao i cht c s sinh ra ch yu duy tr nhng hot ng ti thiu ca cc c quan ni tng, thn kinh v cc chc nng sng ti thiu khc, nng lng ny ch c th gip c th tn ti ch khng th sinh trng. 4.2 4.2.1 Cc yu t nh hng n trao i cht ca c Cc yu t b n trong

+ Loi c : cc loi c khc nhau c cng trao i cht khc nhau. Cc lo i c c tnh n tng t nhau, c no c cng trao i cht ln hn th tc sinh trng ln hn. + Tui c: cng trao i cht ca c nh cao hn ca c ln. C chp c 12 g c cng trao i cht 24,48 kcal/kg.ngy v c chp c 100 g c cng trao i cht 7,97 kcal/kg.ngy + Phi tnh: c c c cng trao i cht cao hn c ci. + Thnh thc sinh dc: trong qu trnh pht trin tuyn sinh dc, cng trao i cht cng khng ging nhau. C giai on IV thnh thc ca tuyn sinh dc c cng trao i cht cao nht. + Dinh dng v i: khi c n no cng trao i cht tng ln v khi b i th cng trao i cht gim xung. C n cc loi thc n khc nhau th cng trao i cht khc nhau. Trong nhng ngy u ca s i, cng trao i cht gim rt nhanh v sau gim rt chm. Khi i nng lng cn thit duy tr s sng ca ng vt hon ton nh vo vt cht ca c th. hu nh th t vt cht trao i l glycogen, lipid v protein.

SLVTS

NVT

110

c, Nagai v Ikeda (1971) c nhn xt ngc li, th t cht trao i l lipid, glycogen v protein. + S vn ng: c vn ng nhiu c cng trao i cht cao hn c t vn ng. 4.2.2 Cc yu t b n ngoi

+ Nhit : khi nhit gia tng th cng trao i cht ca c tng nhng c mt khong nhit , khi nhit tng, cng trao i cht ca c tng khng ng k. Kho ng nhit ny c gi l phm vi nhit thch hp ca c. Phm vi nhit thch hp thay i ty theo loi c v giai on sinh trng ca c. C ln c phm vi thch hp rng hn c nh. i vi mi loi c c mt nhit ti ho cho trao i cht ca n, v nu c cho php mt con c s chn mi trng thun li nht v nhit , v nhit ti ho ny nm trong phm vi nhit thch hp. + nh sng : c c gi di nh sng c cng trao i cht ln hn c gi trong bng ti. + Ma v : c ho t ng mnh nht vo cc thng ma h, km hot ng vo cc thng ma ng; ni cch khc, cng trao i cht ca c cao ma h, thp ma ng do nhu cu nng lng cc thng h cao hn cc thng ma ng. + Dng nc: c nc chy c cng trao i cht cao hn c nc tnh. Lu tc nc gia tng s lm gia tng nng lng c i hi cho vic duy tr mi trng. y khng phi l nng lng bi cn thit m l nng lng gia tng c yu cu duy tr v tr trong mi trng. + CO2 v pH: c sng trong mi trng c nng CO2 cao hay pH thp th c cng trao i cht thp hn c sng trong mi trng c nng CO2 thp v pH cao. C. 1. Dinh dng ca c Khi nim

Dinh dng l s cung cp nhng vt cht cn thit cho vic duy tr i sng (trao i cht). Mt s vt liu s c dng cho s xy dng cc t chc ca c th (ng ha) v mt s cho vic to ra nng lng (d ha). Thc n ca ng vt c th phn chia thnh thc n nng lng v sinh trng nh carbohydrate, lipid v protein v thc n khng nng lng nh cc khong cht, vitamin, nc v oxygen. Phn ln cc thc n khng nng lng ng vai tr b sung cho cc thc n nng lng v mc du rt nhiu trong chng c i hi vi s lng rt nh (cc vitamin, cc nguyn t vi lng) nhng chng rt cn thit cho i sng . 2. Gi tr dinh dng ca khu phn thc n c xc nh bi: - Mt s cn bng gia cc thc n nng lng v sinh trng (carbohydrate, lipid v protein) nhm m bo s cung cp vt liu nng lng cho hot ng ng ha ti ho

SLVTS

NVT

111

v vt liu sng cho s pht trin bao gm kin to cc m, sinh sn v thnh lp cc sn phm m c th cn (hemoglobin, hormone, enzyme, ...). - S hin din ca cc nguyn t (khong cht) nht l nguyn t vi lng v cht xc tc cn thit (vitamin). - S cung cp y thc n ph (nc v oxygen). - Ngoi ra, gi tr dinh dng ca khu phn cn c xc nh bi kh nng c c th tiu ha v hp thu tt nht. N ph thuc trng thi vt l ca thc n v lo i v s lng ca enzyme trong ng tiu ha. 3. 3.1 Nhu cu nng lng Nhu cu nng lng tng cng

Thc n sau khi tri qua qu trnh tiu ha c c th hp thu v s dng vo 2 mc ch: m t phn to thnh vt cht ca c th v mt phn c d ha cung cp nng lng cn thit cho cc hot ng sng. Cc vt cht c th sau cng qua qu trnh d ha to ra nng lng . Nh vy c th ni nng lng thc n hp thu c gm nhng dng nng lng tch ly v nng lng tiu hao. T l gia hai dng ny thay i ty theo loi c v ty theo giai on sinh trng. c pike, t l ny kho ng 1433% calorie hp thu c tch ly trong cc m v khong 6786% nng lng hot ng (Winberg, 1960). c hi (trout) t l ny l 30/70 (Phillips v Brockway, 1959). Ivlev (1939) tm thy c sheatfish 65,7% calorie ca no n hong c tch ly trong c th sau khi hp th non hong hon ton, 34,3% calorie c dng cho nng lng. iu ny s tr nn kh khn gp i khi c sheatfish ln hn do phi tm kim thc n ca chng v t duy tr trong mi trng, l nhng hot ng khng cn thit cho c trong thi gian chng sng trong mt mi trng c bo v v ty thuc vo non hong nh ngun thc n. 3.2 Tnh ton nhu cu nng lng C s dng cc ngun nng lng thc n n vo (C) cho cc mc ch: (1) Tng hp cc m ca c th (P) (2) Cung cp nng lng cho qu trnh trao i cht (R) (3) Sn phm tha c thi ra ngoi (E) C=P+R+E + P c th bao gm cc thnh phn sinh trng (Pg) hoc/v sinh sn (Pr) P = Pg + Pr + Trao i cht tng cng bao gm mt s thnh phn ph - Trao i cht tiu chu n, RS, khi con vt trng thi ngh;

SLVTS

NVT

112

- Trao i cht thng thng (duy tr), RR, khi con vt trng thi hot ng b nh thng; - Trao i cht khi n, RF, ng vi tc ng ng lc c bit (SDA, Specific Dynamic Action) khi con vt va n; - Trao i cht vn ng, RA, khi con vt trng thi vn ng tch cc; R = RS + aRR-S + bRF-S + cRA-S Vi a, b, c l t l thi gian + E c th bao gm phn (F), sn phm bi tit trong nc tiu: urea, ammonia (U) v cht nhy (Muc) + Tng hp cc thnh phn ring bit, ta c C = (Pg + Pr) + (RS + aRR-S + bRF-S + cRA-S) + (F + U + Muc) Theo Brett v Groves (1979) th - C n ng vt: - C n thc vt: 3.3 100C = 29P + 44R + 7U + 20F 100C = 20P + 37R + 2U + 41F

Cch tnh nhu cu thc n hng ngy ca c

Nhu cu nng lng ca c c xc nh gin tip thng qua tiu hao oxygen v thng s h hp RQ (Respiratory quotient) ca c. RQ c xc nh nh sau. Cch tnh RQ: RQ = + i vi trao i cht ng C6H12O6 + 6O2 RQ = VCO2 VO2 = 6CO2 + 6H2O + 637 kcal 6*22,4 6*22,4 = 1 VCO2 VO2

+ i vi trao i cht lipid 2(C55H106O6) + 157O2 RQ = VCO2 VO2 = 106H2O + 110CO2 + 16353 kcal 110*22,4 157*22,4 = 0,7

+ i vi trao i cht protid

SLVTS

NVT

113

Bng thc nghim t chy 200 g protid cn 96,61 L O2 v sinh ra 77,31 L CO2 RQ = VCO2 VO2 = 77,31 L CO2 96,61 L O2 = 0,8

Nh vy RQ ca ng vt bnh thng ch bin ng trong khong 0,71. ng vt n c c khuynh hng c RQ cao hn ng vt n tht. Carbohydrate 4,25 0,82 0,82 1 5,0 14,76 Lipid 9,45 2,03 1,43 0,7 4,7 13,72 Protein 4,25 0,97 0,78 0,8 4,5 13,36 13,60

kcal/g L O2/g L CO2/g RQ kcal/L O2 J/mg O2

(NH3) (Urea)

Cn c vo nhu cu nng lng ca c nui v gi tr nhit ca thc n, ngi ta c th tnh nhu cu thc n hng ngy ca c mt cch gn ng. so snh hiu qu ca cc khu phn thc n ngi ta thng dng h s thc n (Feed Conversion Ratio, FCR) l khi lng thc n cn thit to ra mt n v khi lng tht c. Nhng v gi tr dinh dng ca khu phn ngi ta thng cn c trn s calorie ca thc n cn to ra mt khi lng tht c. Phillips v Brockway (1959) tnh ton trn c brook trout. Loi thc n Thc n t nhin Bt tht kh Tht Nng lng thc n 640 kcal/kg 1540 990 Nng lng 1 kg tht c 2.000 kcal/kg 4.600 2.600 H s thc n 3,1 3 2,6

SLVTS

NVT