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Chng II. SINH L MU


1. H Thng Tun Hon

1.1 Khi nim chung v mu

c, mu l mt t chc lng, mu , vn chuyn trong h thng cc huyt qun.
Mu l thnh phn quan trng nht ca mi trng bn trong c th v m nhn nhiu
chc nng sinh l khc nhau, gp phn iu tit mt cch chnh xc ni mi trng, gi
cho hot ng sng ca c th lun lun bnh thng.

1.2 Hnh thi hc ca h thng tun hon

H thng tun hon (circulatory system) ca c tng t nh cc ng vt c
xng sng khc nhng c nhng khc bit ph hp vi cc iu kin hnh thi, sinh l v
mi trng. C cng c mt h thng bch huyt (lymphatic system) nhng t c bit
n nh cc ng vt c xng sng trn cn.

Bt u t tim ch c mt con ng, ng mch ch bng (ventral aorta), n cc
mang (gills). Tuy nhin, sau khi qu trnh trao i kh xy ra, c nhiu con ng mu
i tr li c th bao gm mt s mao mch nh nhng rt quan trng. ng mch vnh
(coronary artery) ri cung mang th hai (second gill arch) v tr v tim dc theo mt bng
ca ng mch ch bng, cung cp mu bo ha oxygen n tim v n cc nang tuyn
gip (thyroid follicles) phn b ri rc xung quanh ng mch ch bng. T cung mang
th nht (first gill arch) mt mao mch (vessel) chy n mang ph gi (pseudobranch) ri
n tuyn mng trch (choroid gland) nm pha sau mt trc khi ni vi h thng tnh
mch (venous system). Vai tr ca 2 c quan ny c l lin h n s kim sot s thng
kh (ventilation) v trao i kh (gas exchange) vo trong cc dch mt (eye fluids). Tnh
mch mang (branchial vein) hi qui (recurrent) l mt con ng ph (bypass) t cc
mang tr li tim mt cch trc tip v khng phi tt c u ra thuc tim (cardiac output)
cn i vo ng mch ch lng (dorsal aorta) v cc mch mu i ra khc (efferent
vessels). ngha ca tnh mch mang cha c hiu y nhng n c th l phn
quan trng ca u ra thuc tim tr li trc tip tnh mch tim khi c trng thi ngh.

ng mch ch lng l ngun cung cp mu chnh cho cc b phn ca c th. N
cung cp mu cho u (head), cc c thn (trunk muscles), vnh ngc (pectoral girdle),
thn (kidney), v tt c c quan ni tng (visceral organs) l cc mng mao mch chnh
(capillary beds). Sau khi i qua cc mng mao mch chnh, c ba con ng tnh mch
chnh mang mu tr li tim. Cc mao mch phn u tr v tim qua mt i tnh mch
chnh trc (anterior cardinal veins) m s nhp thnh tnh mch chnh chung n (single
common cardinal). Tnh mch chnh chung cng c nhp bi tnh mch chnh sau
(posterior cardinal vein) v mt s tnh mch nh t h thng c phn trc c th
(anterior body musculature). Tuy nhin cc mao mch t h thng c phn sau c th
(posterior body musculature) chy vo tnh mch ui (caudal vein) ri dn vo trong
mng cc mao mch bao quanh cc ng thn (kidney tubules). V y l mng cc mao
mch th cp (second capillary bed) sau khi c cung cp mu bo ha oxygen, n c
xem nh thit k c bit ca h thng ca thn (renal portal system). Cc tnh mch chy
t cc c quan ni tng (ngoi tr gan) to thnh mt h thng ca (portal system) tng
t trong gan (liver). C hai h thng ca dng nh phc v nh cc b (pools) chnh ca
mu m cc sn phm trao i cht (metabolic products) c th khuch tn vo trong

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vi s gia tng rt t v nng ca nhng cht ny. H thng ca thn ch hin din c
v lng c, mc du ng vt xng sng cao ng c mt h thng ca ty (hepatic
portal system).

H thng bch huyt (lymphatic system) t c kho st trn c, nhng hin din
2 dng khc nhau. mt s c, chng hn c hake, ng bch huyt chnh (lymphatic duct)
dng trc (axial) v c nh v trung tm pha trn li thn kinh (nerve cord) bn trong
cung thn kinh (neural arch). c khc, chng hn nhm c hi (salmonids), h thng
bch huyt dng ngoi bin (peripheral) vi mt ng chnh mi knh ng bn (lateral
line canal) v dc theo ng gia lng (dorsal midline). Cng c nhng ng bch huyt
ngn trong mi phin mang (gill filament) c nhim v thu cht dch c lc t bn trong
phin mang v dn chng tr li tim thng qua s ni ca chng vi tnh mch mang gc
mi phin mang. Trong tt c trng hp, cc ng bch huyt phc v em dch m
(tissue fluid) tr li h thng tun hon trong cc m - ni m khng phi tt c cht dch
ny tr li phn h lu (downstream portion) ca cc mng mao mch. Dng nh
khng c bt k cc tuyn (gland) hay hch (node) bch huyt c nh cc ng vt hu
nh.

1.3 Chc nng chung ca h thng tun hon

H thng tun hon phc v cho nhiu chc nng nhng tng qut nht l vn
chuyn, bao gm vn chuyn cc cht kh gia cc m v cc mang v vn chuyn lactate
t cc m n mang v gan v ri vn chuyn glucose tr li cc m. Cc vt cht ngoi lai
c vn chuyn n thn ni m cc thnh phn ha tan c bi tit v thnh phn t
bo b thc bo. S hin din cc cht ny cng dn n sn sinh ra cc khng th c
vn chuyn tr li h thng tun hon. Cc sn phm ca qu trnh tiu ha c vn
chuyn t rut n gan v ri n nhng phn cn li ca c th. Cc t bo mu cng di
chuyn t ni c to thnh n tt c cc phn ca c th. Cc yu t ng mu v
thrombocyte, kt hp bt k v tr tn thng no bt kn vt thng, c a vo h
tun hon m khng ngn cn chnh cc mao mch.


H.1 H thng tun hon ca c

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1.4 Thnh phn ca mu

Mu c hai thnh phn chnh l cc t bo v huyt tng. Cc chc nng ca hai
thnh phn ny i khi phn bit v i khi c chia s bi c hai. Nhng thnh phn ca
huyt tng (hay huyt thanh - huyt tng loi b fibrinogen sau khi lm cho mu
ng) bao gm mt s gii hn cc ion v c v mt thnh phn rng cc hp cht hu c
lin h phn ln n cc chc nng trao i cht. Thnh phn t bo bao gm cc t bo
ring bit c hnh thi v cc chc nng khc nhau.

Hng cu (red blood cell, erythrocyte)
Bch cu (leukocyte)

Huyt cu
Tiu cu (thrombocyte)
Fibrinogen
Nc
Protein huyt thanh
M
ng
Mu
Huyt tng
Huyt thanh
Cht th rn
Mui v c

Cc t bo mu trng thnh c th c xc nh bi hnh thi v cc c trng
bt mu thuc nhum khi quan st di knh hin vi quang hc. Cc t bo hng cu
(erythrocyte hay t bo mu , red blood cell) c nhn, chim u th v s lng v n
nh v kch thc, c s dng nh mt cng c o lng thun tin cho vic tnh ton
kch thc ca cc t bo khc. Ngoi vic o lng kch thc bn ngoi cc t bo,
chng cng c c trng bi t l th tch nhn trn th tch t bo.

Ngoi cc t bo hng cu, cc loi t bo mu chnh khc bao gm bch cu
(leukocyte) v tiu cu (thrombocyte). Cc t bo bch cu tiu biu c nhn tng i ln
v t bo cht t, v cng c phn thnh cc nhm bch cu khng ht (lymphocyte v
monocyte) v bch cu c ht (acidophile, neutrophile v basophile). Cc t bo tiu cu
cha trng thnh trng ging nh cc lymphocyte, v c th dn xut t cc lymphocyte,
nhng thay i trong qu trnh pht trin thnh cc t bo oval. Nhn chung t bo tiu cu
nh hn lymphocyte v c nhn cng nh hn.

Ngun gc ca tt c t bo mu khng c hiu bit y v c th c m t
theo Klontz nh sau. Cc t bo mu c sn xut ra t cc m to mu (hematopoietic
tissue) trong thn v c l trong t tng. Khng c ty xng (bone marrow) hay hch
bch huyt (lymph node) trn c nh ng vt hu nh. Tuy nhin, tn gi cc t bo
mu cng tng t nh hu nh v qu trnh pht trin cng theo mt kiu tng t. Huyt
bo mm (hemocytoblast) l ngun gc ca tt c cc t bo khc. Cc t bo ny c
tng sinh, dn dn bit ha v c hnh thi v chc nng ring bit, thng chng c
pht trin kh r rng trc khi i vo trong mu tun hon. Cc t bo cha trng thnh
ch c th nhn thy trong cc m to mu v s xut hin nhiu cc t bo ny trong mu
tun hon c th biu th cho s hin din ca mt bnh hay yu t bnh l khc.

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H.2 S l thuyt v s to mu c rainbow trout (theo Klontz)

1.5 Lng mu

Tng hp t cc nghin cu cho thy rng lng mu trong c th c t hn so vi
mu ng vt bc cao v nng lng tiu hao cho qu trnh trao i cht ca c t hn.
Lng mu trong c th mt phn tun hon trong tim v mao qun, phn cn li c d
tr trong cc kho cha mu. Lng mu tun hon chim khong 50%; song t l ny lun
lun thay i ty thuc vo trng thi sinh l ca c th: lc bnh thng mu d tr tng
gim bt gnh nng cho tim, khi vn ng th mu d tr i vo h thng tun hon
m bo nhu cu nng lng cho c th. ngi, lng mu l 7,8% so vi trng lng
c th; chim: 7,7%; ch: 6,4-8,2%; th: 5,45%; ln: 4,6%. C nc ngt c lng mu
tng cng chim 2,7% v bin ng trong khong 1,8-4,1%; c bin c lng mu chim
4,1% v dao ng trong khong 1,9-7,3%.

C nhiu yu t nh hng n lng mu trong c th c nh phng thc sng
v trng thi sinh l: c hot ng nhanh nhn c lng mu nhiu hn c t hot ng;
th tch mu gia tng theo tui v giai on thnh thc sinh dc; th tch mu c c cao
hn c ci trng thnh. iu kin sng cng nh hng n lng mu ca c: c tm
Acipenser ruthenus sng sng hoc h c iu kin sng tt (dinh dng tt) th lng
mu nhiu hn so vi nhng c th cng loi c iu kin sng km (dinh dng km).

2. Tnh Cht L Ha Hc v Thnh Phn Ha Hc ca Mu

2.1 Tnh cht l ha hc ca mu

a. Trng lng ring ca mu
Huyt bo mm
(Hemocytoblast)
Nguyn huyt bo nh
(Small lymphoid)
Nguyn huyt bo ln
(Large lymphoid)
T bo bch huyt
(Lymphocyte)
T bo hng cu
(Erythrocyte)
T bo ht
(Granulocyte)
Tiu huyt cu
(Thrombocyte)
Th i thc bo
(Macrophage)
?

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Trng lng ring hay t trng ca mu thay i theo s lng t bo ca n.
Trng lng ring ca mu c nc ngt l 1,035, bin ng trong khong 1,032-1,051;
mu c bin bin ng trong khong 1,022-1,029; ng vt hu nh ni chung khong
1,053 v ngi trong khong 1,05-1,06.

b. Tnh ni ma st ( nht)

Cc nghin cu cho thy nht ca mu c thp hn nhiu so vi ng vt hu
nh (mu c long hn so vi mu ca ng vt hu nh). nht c xc nh bi tr
s ni ma st; v d tr s ni ma st l 3, c ngha l lu tc ca mu chm hn so vi
nc nguyn cht 3 ln. Tr s ni ma st ca mu c ni chung trong khong 1,49-1,83
(ca mu c nhm l 1,70, dao ng trong khong 1,66-2,01); thp hn nhiu so vi ng
vt hu nh, khong 3-6 ( ngi dao ng khong 4-5).

Tnh ni ma st ca mu c quyt nh bi hai yu t: s lng hng cu v hm
lng protein ca huyt tng (plasma protein). Cng mt loi c, khi c cho n tht
tnh ni ma st tng ln, khi c cho n thc vt tnh ni ma st gim xung.

c. p sut thm thu

p sut thm thu (ASTT) ca mu v dch m gi vai tr rt quan trng trong vic
iu ha s trao i nc gia mu v cc m. S thay i ASTT ca ni mi trng s
dn n s thay i c ch trao i nc ca t bo.

p sut thm thu ca mu do cc mui khong trong huyt tng to nn (ch yu
l NaCl). Cc protein trong huyt tng to ra mt phn nh ASTT ca mu, c gi l
p sut th keo.

Cc loi c khc nhau c ASTT khng ging nhau. Nhn chung, c sn c ASTT
ca mu cao hn c xng, c bin c ASTT ca mu cao hn c nc ngt. Cc loi c
sn (bin v nc ngt) v c xng nc ngt c ASTT ca mu cao hn mi trng v
c xng bin c ASTT ca mu thp hn mi trng. Do ASTT ca th dch ni chung v
ca mu ni ring cao hn mi trng bn ngoi nn cc loi c sn v c xng nc
ngt lun lun thi nc tha do thm thu t bn ngoi. Ngc li, do c ASTT ca dch
c th v ca mu thp hn mi trng nn c xng bin phi gi nc v b sung nc
cho c th b p nc b mt do thm thu ra bn ngoi.

p sut thm thu ca mu c tng i n nh. Tuy nhin; trong phm vi khng
nguy hi n c th, n cng thay i theo ASTT ca mi trng. Khi ASTT ca mi
trng tng ln th ASTT ca mu c cng tng ln v ngc li.

- Dung dch sinh l: l nhng dung dch nhn to cha mt s mui v c c ASTT
tng ng vi ASTT ca mu. i vi ng vt hu nh, dung dch sinh l c nng
mui l 0,9%; c l 0,65%.

d. pH

pH ca mu cng l mt ch tiu quan trng phn nh trng thi sinh l ca c th
v s bin ng ca mi trng sng ca c. pH ca mu l thuc vo t l ion H
+
v OH
-


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trong mu. pH mu ni chung l tng i n nh; v d i vi c chp 15
o
C, pH ch
dao ng nh, khong 7,4-7,9. Khi pH mu bin ng thin v acid hoc kim u lm cho
hot tnh ca h thng enzyme trong t bo b nh hng v tnh cht l ha hc ca cc
cht trong t bo cng b thay i, do nh hng n chc nng bnh thng ca c th
mt cch r rt.

Tr s pH trung bnh ca mu c vo khong 7,52-7,71, khng n nh bng ng
vt hu nh. pH ca mu c c th thay i theo pH ca mi trng. Tr s pH ca mu c
bin tng i n nh hn c nc ngt mc d hm lng cc cht m trong mu c
bin t hn c nc ngt l do nc bin c h thng m kh hon chnh nn pH mi
trng tng i n nh trong khi nng ion H
+
trong nc ngt thay i rt ln theo
thi gian v a im. Do , c sng nc ngt thch nghi mnh i vi s thay i pH
ca mi trng hn c sng bin.

Yu t m bo cho s n nh ca pH mu l cc h thng m ca mu. Mt h
thng m gm mt acid yu v mui ca acid vi base mnh. Trong mu c c th tm
thy cc h thng m sau:

H
2
CO
3
* H m bicarbonate:
NaHCO
3

y l h thng m quan trng trong mu v c s lng tng i nhiu.

NaH
2
PO
4

* H m phosphate:
Na
2
HPO
4

HPr

* H m protein:
BPr

Pr: protein, B: Na hay K

y l h m quan trng nht, trong quan trng hn c l h m hemoglobin
(Hb) ca hng cu. Hb c kh nng m gp 10 ln cc protein khc trong huyt tng.

Nh tc dng ca cc h thng m, s cn bng acid/kim lun c gi n nh.
V d, khi acid carbonic sinh ra nhiu s lm pH ca mu gim; khi pH ca mu s
c gip n nh bi h m Hb thng qua phn ng KHb +H
2
CO
3
HHb +KHCO
3
.
HHb c tnh acid yu hn H
2
CO
3
nn khng lm pH mu gim nhiu.

Ngoi ra, s n nh ca pH mu cn c gp phn bi tc ng ca h thng
thn kinh trung ng, c quan h hp v hot ng iu ha ion H
+
ca thn.

- Tc dng ca h hp: khi CO
2
trong mu tng ln, CO
2
kt hp vi nc to
thnh acid carbonic, lm cho pH ca mu gim xung. Mu vi pH thp khi n no kch
thch trung khu h hp ca h thn kinh trung ng hot ng lm tng cng thi CO
2
;

do
gim lng H
2
CO
3
trong mu v pH li c nng ln. Ngc li, nu pH ca mu qu
cao, s c ch trung khu h hp ca thn kinh trung ng, lng CO
2
thi ra ngoi s gim
do tng lng H
2
CO
3
trong mu v pH li c h xung.

- Tc dng ca thn: trong qu trnh trao i cht ca c th, nhiu cht c tnh acid
c sn sinh ra v c a vo mu, chng s kt hp vi kho kim ca mu, nh

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vn duy tr c pH ca mu tng i n nh. Nhng kho kim d tr s dn dn b
hao ht. Thn c tc dng thi i gc acid v gi li gc kim, nh m khi phc kho
kim trong mu.

2.2 Thnh phn ha hc ca mu c

Khi em mu ly tm hoc mu lng t nhin trong mi trng lnh s din ra
qu trnh phn chia thnh phn dch lng gi l huyt tng (cht dch c mu vng nht
hoc khng mu) v phn lng ng (c mu ) c hnh thnh t cc thnh phn hu
hnh ca mu (hng cu, bch cu, tiu cu). Huyt thanh l huyt tng b loi
fibrinogen.

T l gia th tch huyt cu v huyt tng thay i theo ging loi v phng
thc sinh sng ca c. Thng thng t l th tch huyt cu so vi mu (hematocrite)
chim khong 27%, cao nht l 36% nh c chp (Cyprinus carpio), thp nht l 16%
nh c hm ch (Lophius piscatorius).

a. Nc

Nc l thnh phn c t l ln nht trong mu, chim ti 80%. Trong huyt tng,
nc chim ti 90-92%, hm lng nc trong hng cu t hn 65-68%. Khi b mt nc
nhiu s lm mu c qunh li, qu trnh trao i cht s ngng tr.

Nhn chung nc trong mu c xng t hn c sn, c con nhiu hn c trng
thnh.

b. Protein

Protein l thnh phn ch yu trong cht kh ca huyt tng. Cc nghin cu cho
thy rng protein trong mu c bin ng rt ln. S bin ng ny khng ch din ra trong
cc loi khc nhau m ngay c trong cng mt loi v thm ch trong cng mt c th.

Trong thnh phn protein ca mu c 3 nhm chnh: albumin, globulin v
fibrinogen.

+Fibrinogen: sinh ra gan, tin cht ca fibrin (si huyt), c vai tr ng mu.

+Albumin: sinh ra gan, lin kt vi cc lipid, hormone. p sut thm thu huyt
tng phn ln l do albumin.

+Globulin: l cht vn chuyn cc lipid v steroid, st v ng. Khng th l mt
phn ca globulin.

Hmlng protein trong huyt thanh ca c thay i trong khong 2,5-7 mg%,
trong khi hm lng protein mu ngi thay i trong khong 7,5-8,5 mg%; cho thy
lng protein trong huyt thanh trong mu c thp hn ngi. Mt vi nghin cu cho
thy lng protein trong huyt thanh thay i ph thuc vo iu kin dinh dng ca c.
V d: c chp c nui trong ao c thc n t nhin phong ph th lng protein trong
mu l cao hn c chp c nui mt phn bng thc n t nhin v nhn to.


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Hm lng protein trong mu c cn thay i theo ma v. V d: c chp 1 tui
sng vng n i qua ma ng protein huyt thanh gim t 3,8 mg% cn 2,7 mg%,
albumin hu nh mt ht. Qua mt thi gian bt mi bnh thng, hm lng protein huyt
thanh dn dn c khi phc.

Protein trong mu c cc vai tr sau y:

- Duy tr p sut thm thu cho mu, cn gi l p sut th keo;
- Tham gia vo h m ca mu (Hb);
- ng vai tr quan trng trong qu trnh ng mu (fibrinogen);
- L nhng khng th bo v c th: globulin, khng th chng li vi trng v
virus.

Protein huyt tng trong c th lun lun b phn gii v khng ngng c tng
hp v trc tip tham gia vo qu trnh trao i cht ca c th.

c. Nit phi protein (non-protein nitrogen, NPN)

Nit phi protein l nhng sn phm trung gian v sn phm cui cng ca qu trnh
trao i cht protein. Khi tch cc protein trong huyt tng ca c th cn li mt s hp
cht cha gc nitrogen. Cc nghin cu cho thy rng s lng cc cht cha nitrogen
trong mu c kh cao. V d: c chnh Nht Bn (Anguilla japonicus) vo ma h trong
mu cha 125,6 mg% hp cht cha nitrogen.

S lng cc cht cha nitrogen gia tng khi cc hot ng bin dng ca c gia
tng. Ngi ta cn pht hin ra rng c xng s lng hp cht ny thp hn nhiu ln
so vi c sn.

- Ammonia (NH
3
): l mt vt cht c c nng thp trong mu phn ln ng
vt. Nng ammonia trong mu c cao hn ng vt hu nh nhng nh hn 0,1 mg%.
Phn ln c xng nc ngt l ammonotelic (ng vt bi tit amn). Vai tr ca s bi
tit NH
3
l cn bng acidbase mu.

- Urea (CO(NH
2
)
2
): l mt cht t c, n c to thnh t NH
3
v ha tan trong
nc nhiu hn ammonia. C sn c gi l ureotelic (ng vt bi tit urea). Mt s c
xng cng bi tit lng ln urea. Cc loi c bin c nng urea mu 22,5 mg%, cao
hn cc loi c nc ngt 1 mg%. S d cc loi c sn bin c nng urea mu cao l
duy tr p sut thm thu ca mu cao hn mi trng ca chng.

- TMAO (Trimethylamine oxide): l mt cht ha tan khng c. mt s c bin
n c bi tit chim ln 1/3 lng nit ca chng. TMAO c bin cao hn c nc
ngt. C hi (salmon) v chnh (eel) khi bin c TMAO cao hn khi nc ngt. C
Spiny dogfish (Squalus acanthias) c hm lng TMAO mu tng i cao v trn 90%
TMAO c lc bi thn c ti hp thu gi nc c hiu qu.

d. Glucid

Glucid l thnh phn hu c ch yu trong huyt tng. ng trong huyt tng
ch yu dng glucose. Hm lng ng ng vt mu nng bin i trong phm vi
hp, ngi t l ny khong 0,10,12% khi lng mu nu vt qu 0,18% th glucose

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s b o thi theo ng nc tiu v ngi ta mc bnh tiu ng. Nhng c th hm
lng ng bin i kh ln. Hm lng ng trong mu c sn thp. Hm lng
ng trong mu c xng bin c lin h n tp tnh sng ca c: c hot ng chm
chp c lng ng huyt thp, c hot ng mnh c lng ng huyt cao. Khi lng
ng huyt ca c hot ng tng i mnh gim xung th n hot ng chm chp,
cn loi c c tp tnh hot ng chm chp th nh hng khng r rt. cc loi c
nc ngt, s quan h gia lng ng mu v tp tnh hot ng khng r rt nhng c
s khc nhau gia cc ging loi. V d: c chp (C. carpio) c lng ng mu l 58
145 mg%, cn c vn (Abramis brama) l 122230 mg%.

Hm lng ng trong mu c thay i ty theo trng thi sinh l ca c nh c
tng vn ng th lng ng trong mu tng ln. iu kin mi trng ngoi khng
thun li nh thiu oxygen, chn thng do hot ng c hc, dn p c trong khi nc
cht hp, v.v. cng lm tng lng ng trong mu. Mt s nghin cu cho thy hm
lng ng trong mu c ph thuc vo gii tnh (c c cao hn c ci) v s thnh
thc sinh dc ca c.

e. Lipid

Trong huyt tng lipid khng dng t do m kt hp vi protein thnh mt hp
cht ha tan. Mt trong cc cht lipid c nghin cu nhiu l cholesterol. Cc nghin
cu cho thy rng trong mu c chnh (eel) hm lng cholesterol t trn 700 mg%, hm
lng c nhm (shark) l 21 mg%. Khi lu gi c trong iu kin nhn to cc b xi
mng v tuyn sinh dc ca c ang trong giai on thoi ha th ng thi lng
cholesterol trong mu c gia tng ng k. Trong qu trnh chn ca tuyn sinh dc hm
lng cholesterol trong mu gim thp rt nhiu. iu ny khng nh khi hm lng
cholesterol trong mu cc c th trng thnh gia tng, chnh l du hiu ca s
thoi ha tuyn sinh dc.

f. Cc cht v c

Trong mu c gm c mt s cation ch yu nh Na
+
, K
+
, Ca
2+
, Mg
2+
v thng
kt hp vi mt s anion nh Cl
-
, CO
3
2-
, PO
4
3-
, trong mui NaCl chim n 8695%.
Thnh phn v t l cc mui v c trong mu c cng tng t nh mu ca ng vt
hu nh, ng thi cng tng t nh thnh phn v t l ca mui trong nc bin.

Mui trong mu c l thnh phn to nn nng thm thu ca mu. Cc ion K
+
,
Na
+
cn cho s hng phn ca h thn kinh, co bp c, nht l c tim; Ca
2+
cn cho vic
to xng cng nh trong qu trnh ng mu. S lng cc mui v c tng cng thay
i t 1,31,8% khi lng mu. Hm lng v t l mui trong mu ca cc loi c khc
nhau th khc nhau, gia c c v c ci ca cng mt loi cng khng ging nhau v
thay i theo chu k i sng v trng thi sinh l c th.

3. Thnh Phn Hu Hnh ca Mu (Cc T Bo Mu)

Cc thnh phn hu hnh ca mu bao gm: hng cu (erythrocyte), bch cu
(leucocyte), tiu cu (thrombocyte).

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15

3.1 Hng cu

a. Hnh thi, kch thc, s lng v chc nng

. Hnh thi

Hng cu l loi huyt cu c s lng nhiu nht trong cc t bo mu. Hng cu
c trng thnh phn ln hnh bu dc. Hng cu ca c c nhn, hai mt li ra (tng
t hng cu ca chim, b st v lng c nhng khc hng cu ca ng vt hu nh hnh
trn dp, khng nhn v c 2 mt lm vo). Do c nhn nn hng cu ca c c cng
trao i cht cao, tiu hao lng oxygen ln.

. Kch thc

c biu th a*b, n v tnh l
a: ng knh nh
b: ng knh ln

- Ty tng ging loi c khc nhau m kch thc hng cu khc nhau
- Kch thc hng cu cng thay i theo tui c
- ng vt tin ha cng cao th kch thc hng cu cng nh

. S lng

S lng hng cu thng c m trong 1 mm
3
mu, n v tnh l triu t
bo/mm
3
. Trong iu kin bnh thng, s lng hng cu ca mi loi c l n nh, n
phn nh tp tnh sng vtnh n ca c: c sng tng mt c s lng hng cu thp, tng
dn c sng tng gia v cao nht c sng tng y.


H.3 Hnh dng cc t bo mu

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16
Cc nghin cu cho thy s lng hng cu trong mu c thay i rt ln. c
nc ngt thay i t 13,5 triu TB/mm
3
. c bin t 0,94 triu TB/mm
3
. S lng
hng cu c tng quan cht ch vi cc hot ng ca c (c bi li nhanh, hot ng
mnh s lng hng cu cao). S lng hng cu trong mu c gia tng theo tui c trong
cng mt loi. S lng hng cu thay i ph thuc vo gii tnh, con c s lng
hng cu thng cao hn con ci do c c c cng trao i cht cao hn c ci.
Trong thi k thnh thc tuyn sinh dc s lng hng cu gia tng v khi sinh sn s
lng hng cu gim thp. Sau khi kt thc ma sinh sn th s lng ny li gia tng.
Trong iu kin thoi ha ca tuyn sinh dc, s lng hng cu trong mu gim thp.

S lng hng cu trong mu thay i theo ma trong nm. V d: c chp (C.
carpio) vo ma ng th s lng hng cu gim cn 2 triu TB/mm
3
; ngc li, vo ma
xun s lng hng cu tng ln 2,85 triu TB/mm
3
.

Trong iu kin c khng c cho n mt thi gian di th s lng hng cu s
gim thp. Ngoi ra, s lng hng cu cn thay i rt mnh ph thuc vo cht lng
thc n c cung cp.

Khi oxygen trong mi trng nc gim xung t ngt th s lng hng cu ca
c tng ln mnh lit, nhng nu c sng lu di trong mi trng thng xuyn thiu
oxygen th s lng hng cu li gim xung, khi a c tr li mi trng bnh thng th
s lng hng cu li dn dn khi phc. Khi hm lng CO
2
t do tch ly nhiu trong
nc cng lm tng s lng hng cu.

pH trong nc t nhiu cng lm thay i s lng hng cu ca mu. C sng
trong mi trng pH thp (acid) th s lng hng cu trong mu cao hn mi trng pH
cao (kim tnh). S thay i nhiu ca p lc nc cng dn n s thay i ln v s
lng hng cu.

Nhng trng thi bnh l ca c th do cc loi k sinh trng gy nn, ni chung
cng lm thay i s lng hng cu.

. Chc nng

Chc nng ch yu ca hng cu l vn chuyn O
2
t mang ti cc m v gp phn
vn chuyn CO
2
t cc m n mang (chc nng h hp), tham gia duy tr thnh phn cc
ion ca mu, iu ha pH mu. Mng hng cu c tnh n hi do d thay i hnh
dng; c tnh ny gip hng cu theo mu qua c nhng mao mch c kch thc nh
hn kch thc hng cu.

c bit, trong hng cu c huyt cu t Hb, cht vn chuyn kh O
2
v CO
2
. Tuy
nhin, ch khi Hb tn ti bn trong hng cu th mi c tc dng; khi hng cu v ra, Hb i
vo huyt tng s b phn gii ha hc nhanh chng.

b. Huyt cu t Hemoglobin (Hb)

Hb chim 90% trng lng cht kh ca hng cu v lm cho hng cu c mu .
Hb l mt protein phc tp gm mt phn t globin (96%) kt hp vi 4 phn t Heme
(4%). Globin l mt protein nn mang bn cht c trng cho tng loi.


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St bnh thng dng Fe
2+
. Chc nng ca Hb l mt cht mang oxygen ty
thuc vo s lin kt lng lo ca Fe
2+
vi oxygen. Khi lin kt vi oxygen, ha tr ca st
khng thay i. St c th b xi-ha thnh Fe
3+
bi cht xi-ha mnh thnh MetHb v
Hb lc ny khng cn c th kt hp thun nghch vi oxygen. Oxygen kt hp vi Hb
theo t l mt phn t O
2
vi mi nguyn t st. CO (oxit cacbon) cng c th kt hp
thun nghch vi Hb nhng lc lin kt ca CO thng ln hn nhiu so vi oxygen (gp
250 ln).

Hm lng Hb thng c biu th bng % hay s gr Hb c trong 100 mL mu
(g%). Hm lng Hb ca cc loi c sn tng i thp: 1,7-5,8 g%, ch bng 1/2 hoc 1/3
c xng 414,7 g% (Stroganov, 1962). Hm lng Hb ca c xng bin c lin quan
n tp tnh vn ng ca c: nhng c hot ng nhiu c hm lng Hb cao, nhng loi
c t hot ng v sng y c hm lng Hb thp. Quan h gia hm lng Hb v tnh
hot ng ca c nc ngt khng r rng. Tuy nhin; hm lng Hb ca c nc ngt
biu hin s khc nhau r rt theo phi tnh: c c lun c hm lng Hb cao hn c ci.
C trng thnh c hm lng Hb cao hn c nh.

C sng vng nc thiu oxygen th c lng Hb cao hn c sng vng giu
oxygen, c c c quan h hp ph hp th c oxygen trong khng kh c hm lng Hb
cao hn c ch ly oxygen trong nc.

Ngoi ra hm lng Hb cn lin quan ti thnh thc ca tuyn sinh dc. Khi h
s thnh thc sinh dc (t l gia khi lng tuyn sinh dc v khi lng c) ca c chp
hoang di (C. carpio) tng t 5 ln 15% th hm lng ca Hb t 41,8% tng dn ln n
43,5%; khi h s thnh thc sinh dc tng ln ti 17% th hm lng Hb cng tng mnh
v t ti 51,5%.

c. Tc lng ca hng cu

Do hng cu c t trng ln hn huyt tng nn khi dng mt cht khng ng (v
d: mui citrat) lm cho mu khng ng v cho mu vo ng nghim ng. Sau mt
thi gian cc hng cu s lng xung v pha trn ng nghim l mt lp huyt tng trong
sut. cao ca ct huyt tng biu th cho tc lng ca hng cu. Tc lng hng
cu ca cc loi c khc nhau: c hi ch (Oncorhynchus keta) tui 1
+
l <1 mm/gi,
c chp (C. carpio) v c dic (Carassius carassius) tui 1
+
l 23 mm/gi. Tc lng
hng cu ca c c nh hn c ci, v tc lng hng cu ca c c v c ci u tng
ln trong thi k thnh thc tuyn sinh dc.

d. Sc khng ca hng cu

Trong iu kin sng bnh thng, p sut thm thu bn trong hng cu bng vi
p sut thm thu ca huyt tng nn hng cu gi c hnh dng v kch thc nht
nh. Nu cho hng cu vo dung dch u trng (ASTT cao) th hng cu s b mt nc
teo li; ngc li nu cho hng cu vo dung dch nhc trng (ASTT thp) th hng cu
s ht nc trng to ra. Nu trng to qu mc hng cu s v ra cc cht bn trong hng
cu s tan vo trong dung dch. Hin tng ny gi l s tan mu.

trc nh sc khng ca hng cu c th dng cc dung dch NaCl nhc
trng vi cc nng khc nhau. Cho vo cc dung dch NaCl mt git mu, yn mt
thi gian ghi nhn nng NaCl lc bt u xut hin hin tng tan mu sc khng

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hng cu nh nht, v nng NaCl lc bt u tan mu hon ton sc khng hng
cu ln nht.

Sc khng ca hng cu khc nhau cc loi c. Dung dch ng tng ca
mu c tench (Tinca tinca) l dung dch NaCl 0,83% v sc khng hng cu nh nht
dung dch NaCl 0,41%. c tm (sturgeon), sc khng HC nh nht l 0,38% v ln
nht l dung dch NaCl 0,20%. c chp, sc khng HC nh nht dung dch NaCl
0,27% v ln nht 0,24%. Cng nh cc ng vt cao ng, tnh bn vng v sc
khng hng cu ca c khng phi l mt hng s, n thay i di nh hng ca cc yu
t ngoi cnh thng qua h thn kinh giao cm lm thay i thnh phn ha hc ca mu.
V d: khi c t nhin t nc lnh di c vo nc nng hoc ngc li, s kch thch ca
s thay i nhit i vi c quan cm gic ngoi da s lm gim tnh bn vng ca
ASTT ca hng cu c.

3.2 Bch cu

a. Cc loi bch cu, s lng, chc nng

. Cc loi bch cu

Bch cu c phn bit bng cc tiu chun v kch thc, hnh dng, cu trc ca
nhn v cc ht bt mu thuc nhum trong t bo cht.

+Bch cu khng ht: t bo cht khng c ht bt mu thuc nhum, gm BC n
nhn (monocyte) v t bo lympho (lymphocyte). Bch cu khng ht ph bin l
lymphocyte c kch thc nh, nhn trn to v t bo cht t. Nhn khng chia thnh nhiu
thy, kch thc nh hn rt nhiu so vi t bo bch cu c ht.

+Bch cu c ht: c trng ca cc t bo ny l t bo cht c nhiu ht bt mu
thuc nhum, nhn chia thnh nhiu thy. nhm ny c th chia ra cc t bo nh sau:

- Bch cu c ht a acid (acidophyle, eosinophyle)
- Bch cu trung tnh (neutrophyle)
- Bch cu c ht a base (basophyle)

c cng gp c hai loi bch cu khng ht v bch cu c ht. Bch cu c ht
thng rt him, trong BC trung tnh v a acid thng thy nht, cn bch cu a base
hu nh t gp.

. S lng bch cu

Bch cu trong mu c c s lng t hn hng cu khong t 10100 ln nhng
nhiu hn ng vt hu nh. S lng bch cu thay i t loi ny sang loi khc. V d:
s lng BC th l 8.000 TB/mm
3
, g l 30.000 TB/mm
3
, c m trng l 51.000
TB/mm
3
, ngi l 6.0008.000 TB/mm
3
.

. Chc nng ca bch cu

+Chc nng bo v c th


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Nhim v chnh ca bch cu l chng li s xm nhp ca vi khun vo c th.
Bch cu trung tnh c kh nng thc bo cc vt l c kch thc nh nh vi khun v c
kh nng di chuyn xuyn qua cc mao mch, c th chuyn ng nh hng n nhng
ni b vim nhim.

Bch cu a acid c kh nng lm mt c t ca vi khun v cc protein l, kh
nng thc bo yu.

Bch cu a base hin din vi mt t l thp trong mu, khng c kh nng vn
ng v thc bo.

Bch cu khng ht monocyte c kh nng thc bo. Cc lymphocyte l nhng t
bo c kh nng min dch. Cc lymphocyte sn xut ra cc khng th -globulin v nht
l -globulin, y l mt khng th chng vi trng rt mnh.

+Cc chc nng min dch

Tri vi ngh rng c xng khng c mt p ng min dch (immune reponse),
ngi ta tm thy mt p ng min dch vi mt khng nguyn (antigen) xy ra trn
cc c hi (salmonids) mc du chm hn so vi ng vt hu nh v phn ln xy ra
gii hn nhit trung bnh hay cao.

Vic sn xut mt khng th l mt qu trnh xy ra trong lymphocyte nh s p
ng i vi s hin din ca vt cht protein ngoi lai bao gm c t bo nguyn vn,
chng hn vi trng. Cc lymphocyte c nhy cm bi protein ngoi lai (khng nguyn)
v nu chng sng lu, s phng thch mt vt cht protein mi (khng th, antibody)
vo trong huyt tng trung ha protein ngoi lai. Cc protein hay t bo ngoi lai c
bao bc vi cc phn t khng th v trung ha cc kh nng c ca chng v cho php
cc t bo thc bo (phagocytic cell) nut chng mt cch an ton. Cc khng th thng
c nhiu hn mt v tr phn ng v nh th c th lm cho cc phn t v t bo ngoi lai
nh vo cng mt khng th kt dnh li vi nhau (agglutinate). Cc phn t khng th vi
mt u nh vo mt t bo vi trng cng c th hot ha mt protein khng phi l
khng th (non-antibody) c gi l th b sung (complement). Cc phn t khng th v
b sung c th phn gii (rupturing) cc t bo vi trng. c, th b sung c sn xut t
cc protein globulin trong mu v khng chuyn bit (hot ng vi bt k khng th no).
Vic sn xut khng th xy ra phn ln t tng v u thn, gan v mu ngoi bin c
th sn xut khng th vi mc t hn, v mt cch c bn, bt c ni no c s tp
trung lymphocyte.

Vic sn xut ra khng th (p ng min dch) thng l mt phn ca qu trnh
vim nhim. ngi, cc du hiu bn ngoi ca s vim nhim bao gm au, sng, v
st. c khng th o lng s au, st v du hiu . Nh vy du hiu sng ty l biu
hin ca s vim nhim. Tuy nhin mc t bo, c s tng t gia c v ng vt
hu nh. Histamin v cc vt cht khc c gii phng v tr vim nhim gy ra s co
mch (vasoconstriction) chn dng mu i vo vng vim nhim v ngn s lan rng
ca cc mm bnh. iu ny gy ra s sng ty. Mt mng li fibrin cng c to thnh
xung quanh v tr tn thng. S vim nhim da c cng kch thch s phn tn t bo
melanin (melanocyte) lm da tr nn en v nh vy che lp du hiu ca vng tn
thng. Sau khi vng vim nhim c bao vy, cc t bo thc bo s i vo v tiu ha

SLVTS NVT
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vt cht ngoi lai ri i tr li thn v t tng trong thi gian bnh khong 2 tun cho n
khi mt khng th thc s c gii phng vo trong mu.

Sinh vt gy bnh s khng th ng trong qu trnh vim nhim ny. Nhiu vi
trng sn sinh ra c t git cc lymphocyte v cc t bo khc. Cc vi trng cng sn sinh
ra cc tc nhn phn gii (lytic agents) lm ha tan vch fibrin quanh vng vim nhim v
xm nhp tr li dng mu v cc khng gian c th khc. Khi t c s xm nhp ny,
cc sinh vt gy bnh nh Vibrio angullarium v IHN virus (infectious hematopoietic
necrosis) c th lm cht cc c hi (salmonids) trong vng 5 v 9 ngy mt cch tng
ng. Thi gian gy cht ny th ngn hn thi gian cn thit cho c sn sinh ra cc khng
th. Nh vy p ng min dch ch c ch cho c khi chng tn ti sau s sc ban u
vi mt s ly nhim.

C khng phi hon ton khng c bo v khi s tn cng ban u ca sinh vt
gy bnh. Trc ht, vy (scale) v cht nhy (mucus) hot ng nh ro cn c hc v
ha hc (mechanical and chemical barriers) chng li s xm nhp. mt s loi c,
dng nh c cc cht dit khun v nm (bactericide and fungicide) trong cht nhy ca
chng. Cc khng th c to ra trong p ng min dchi khi cng xut hin trong
cht nhy. Mt s loi c c s khng t nhin r rng i vi cc bnh lin h gn gi
vi loi. Phn ln cc ng vt c xng sng c cc cht ngng kt t nhin (agglutinin)
c kh nng kt dnh nhiu loi khng nguyn. Nhm c hi salmonid cng c interferon
tc nhn khng virus chung trong mu ca chng. ng xm nhp qua ng tiu ha ca
vi trng th kh khn v pH thp trong d dy v hot ng phn gii protein ca cc
enzyme trong d dy v rut. Tuy nhin khi c tnh trng stress, rut b tc (khng c
hot ng nhu ng, peristalsis) dn n s ln men ym kh cc cht cha trong rut,
cng lc cc enzyme trong rut c th tn cng vch rut. iu ny cho php s tng sinh
nhanh chng ca nhiu tc nhn gy bnh cng nh s xm nhp d dng ca chng vo
trong tun hon mu. Ni chung, i vi phn ln c u c t hay nhiu cc sinh vt gy
bnh tim nng chung quanh, trn v trong c th v ch i mt trong nhng ro cn m
ra tn cng.

S to min dch th ng (passive immunization) l gii thiu nhng khng th
c to ra trc (trong c khc hay trong ng vt hu nh) vo trong c th c nhn
to ra s min dch th ng tc thi (passive immunity). S to min dch tch cc (active
immunization) l gii thiu sinh vt gy bnh c x l trc lm mt kh nng
gy bnh ca chng nhm kch thch mt c nhn to ra s min dch tch cc (active
immunity) ca ring n.

Mt vi con ng gii thiu vaccine c th vi nhng mc thnh cng
khc nhau. Mt trong nhng con ng kh ph bin l tim chng. Vic tim vaccine vo
xoang thn (interperitoneal), di da (subcutaneous) v trong c (intramuscular) cho hiu
qu km. Vic tim vo khng gian gia cc c trn trc v cc gai thn kinh ct sng
ng gia lng cho hiu qu cao hn, c l do vic tip nhn vt cht min dch qua ng
bch huyt lng (dorsal lymphatic duct). Vic chng nga qua con ng ming bng cch
trn vaccine vo thc n cho kt qu thay i, nht l khi c b bnh thng gim hay
ngng n. Nh vy vic to min dch qua con ng ming c tc dng phng hn l tr
bnh. Mt phng php chng nga khc ang c th nghim nhng kh trin vng
l phng php vn chuyn u thm thu (hyperosmotic delivery). Trc ht ngm c vo
dung dch mui v urea mnh (5,3%) trong 2 pht v sau vo trong dung dch vaccine.
Con ng xm nhp ( c hi salmon) qua ng bch huyt ngay di (hay c l c ni

SLVTS NVT
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vi) knh ng bn (lateral line canal). Nhiu phng php chng nga gn y ch n
gin l nhng c vo dung dch vaccine m khng cn dung dch u thm thu cng cho
kt qu chp nhn c.

3.3 Tiu cu (thrombocyte)

Tiu cu l nhng t bo nh, nhn chim ch yu th tch t bo. Cho n nay
ngi ta ch c th phn bit c tiu cu v bch cu lympho (lymphocyte) di knh
hin vi in t hoc thng qua cc phn ng min dch. Bng knh hin vi quang hc rt
kh khn phn bit 2 loi t bo ny trong mu c.

Chc nng chnh ca tiu cu l gii phng cht thromboplastin (thrombokinase)
gy ng mu. Tiu cu cn c c tnh kt dnh nh vy m gp phn ng ming cc vt
thng li.

4. C Ch ng Mu

Khi t chc ca c th c b tn thng, mu s chy ra ngoi. Sau mt thi gian
khng lu th ng c li, kt thnh mt khi mu lp kn vt thng lm cho mu ngng
chy ra. Qu trnh phn ng sinh l ny c lin quan ti s chuyn bin ca fibrinogen
trong mu thnh fibrin.

C 3 thnh phn chnh i vi h thng ng mu: (1) mt lot cc enzyme xc tc
cc phn ng dn n vic sn xut ra si fibrin, (2) cc t bo mu tr nn dnh v to
thnh mt khi v (3) h thng enzyme phn gii fibrin (fibrinolytic) dn n ha tan cc
mu ng (clot). H thng ng mu hot ng trong theo mt kiu ngn chn s r r
trong cc mch mu m khng ngn mu chy trong cc mch mu nh hn bng cch
iu chnh s cn bng gia 2 thnh phn i lp ny.

Fibrinogen ha tan trong huyt tng c chuyn i thnh fibrin dng si khng
ha tan qua tc dng ca enzyme thrombin. Thrombin khng hin din trong mu m c
hnh thnh t prothrombin, y l mt protein ca huyt tng c tng hp t gan.
Thrombin c hnh thnh qua phn ng gia prothrombin v thromboplastin
(thrombokinase, mt sn phm ca thrombocyte hoc cc m b thng tch).

C ng u vi 2 tr ngi lm cho qu trnh ng mu tr nn kh khn hn so
vi ng vt trn cn. Bt k vt thng lm mu chy trn bn mt c th c mt s ng
kh khn v thi gian v cc enzyme cn thit v cc thnh phn ng mu b pha long
hoc ra tri trc khi cc mu ng c to thnh. Tuy nhin c mt thun li cho c l
chng c mt h thng ng mu hot ng rt nhanh ti thiu s pha long ny. Mu
c c th ng c trong vng 20-30 giy so vi mu ngi l khong 7-8 pht. Nhn
chung, tnh trng stress lm gim thi gian ng mu v tng cng ng mu.

C thc nhanh h thng ng mu ca chng bi mt s gia tng s lng cc
thrombocyte trong mu tun hon. Thrombocyte khng hin din ph bin trong mu ca
cc c hi khc nhau khi khng b quy ry nhng c gia tng nhanh chng v s lng
trong v sau mt tnh trng stress (stressful situation). Dng nh cc thrombocyte tng
ln ny c phng thch t mt ni d tr (c l l t tng), nhng khng r l nhng t
bo ny ang c s dng v thay th bng nhng t bo mi hay s tr v kho d tr
c s dng li.

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Cc nghin cu cho thy rng c ch ng mu c ging nh ng vt hu nh
nhng c mt s khc bit v chi tit. V d, thi gian ng mu ca ngi c thc y
nhanh di iu kin stress khong 30% v mt s gia tng tng t s lng cc tiu
huyt cu (platelet) nhng thi gian ng mu ca c c th tng nhanh gp 3-5 ln. Hn
na t bo ng mu ca hu nh (tiu huyt cu) khng c nhn, tui th ngn v c th
b ph hy; trong khi thrombocyte ca c c nhn, tui th c th di hn v c th ti s
dng mc du c hai thc hin cc chc nng ging nhau.

Fibrinogen (ha tan)

Prothrombin Thrombin
(tng hp t gan) Ca
2+
(enzyme)
Fibrin (dng si)
Thromboplastin
(T thrombocyte hay
cc m b tn thng) Mu long Mu ng


Cc mu co Huyt thanh

H.4 S c ch ng mu