Sinh Thai Bien Va Ven Bo

TRONG AI HC THY LI - VIN NGHIN CU THY LUC DELFT
BI GING
SINH THI BIN V VEN B

Son tho : PGS. TS. L nh Thnh
Tr gip : TS. Jeroen Wijsman, TS. Mindert de Vries

ti liu tham kho

1. Chng trnh khoa hc v cng ngh cp Nh nc KT-02, Bo v mi trng v Pht
trin bn vng, H Ni 1995.
2. L Din Dc, Kim k t ngp nc Vit Nam, H Ni 1991.
3. Nguyn Ngc Anh, nh gi tc ng ca hai cng trnh Du Ting v Tr An n
tnh hnh xm nhp mn h lu.
4. Ngn hng Th gii, Ngn hng Pht trin Chu A, FAO, UNDP, Vit Nam- nh
gi tng quan v ngnh Thy li, H Ni 1996.
5. V Trung Tng, C s sinh thi hc, NXB Gio dc, 2000
6. Nguyn Vn Tuyn, Sinh thi v Mi trng, NXB Gio dc, 2000
7. Phng Ngc nh, Ti nguyn bin ng Vit Nam, NXB Gio duc, 1999
8. L nh Thnh, Environmental and Ecological Aspects in Estuary and Coastal Areas
of Vietnam, Workshop in Upgrading the training capacity on coastal engineering at
HWRU, 1999.
9. Asian Development Bank, Coastal and Marine Environmental Management for Ha
Long Bay, SRVN, 1995
10. J.L Chapman, M.J Reiss, Ecology: Principles and Applications, Cambridge
University Press, 2001
11. Jeffrey S. Levinton, Marine Biology: Function, Biodiversity, Ecology, Second
Edition, Oxford University Press, 2001
12. Carol M. Lalli, Timothy R. Parsons, Biology Oceanography, Pergamon Press, 1996
13. Gilbert Barnabe and Regine Barnabe Quet, Ecology and Management of Coastal
Water: The Aquatic Environment, Praxis Publishing, 2000
14. James Perry, Elizabeth Vanderklein, Water Quality: Management of a Natural
Resource, 1996
15. Ton That Phap, Management of Biological Resources in Tamgiang lagoon from a
socio-economic cultural Viewpoint, 2000
16. FAO- Regional Wood Energy DevelopmentProgramme in Asia, Mangrove
production and protection: A changing resource system: Case study in Can Gio
district, Vietnam, 1993.
17. Australian Institue of Marine Science and others, Project PN 9412: Mixed Shrimp
Farming Mangrove Forestry Models in the Me Kong Delta, 1999.

ii
Mc lc
gii thiu
phn 1 khi qut chung v sinh thi
Chng 1: Tng quan v sinh thi hc
1.1 Gii thiu. ................................................................................................... 3
1.3 Cc nhn t mi trng................................................................................ 9
1.4 Ni sng.... ................................................................................................ 15
1.5 S thch nghi.............................................................................................. 18
1.6 Cu hi v tho lun .................................................................................. 18
Chng 2: Qun th v qun x
2.1 ng lc hc qun th............................................................................... 19
2.2 S tng tc gia cc loi ......................................................................... 26
2.3 Bc dinh dng.......................................................................................... 30
2.4 Qun x.... ................................................................................................. 36
phn 2 sinh thi bin v ven b
Chng 3: Mi trng bin
3.1 Gii thiu chung........................................................................................ 42
3.2 Cc c trng v sinh ca nc bin.......................................................... 55
3.3 Cu hi tho lun....................................................................................... 68
Chng 4: Thc vt ph du v nng sut s cp
4.1 S phn loi ............................................................................................... 69
4.2 Cc phng php thu thp mu.................................................................. 74
4.3 Sn phm s cp ........................................................................................ 76
4.4 Cu hi tho lun
Chng 5: ng vt ph du
5.1 Phng php phn loi............................................................................... 90
5.2 Cc phng php thu thp mu................................................................ 100
5.3 Phn b theo ng vt ph du theo chiu su........................................... 101
5.4 Thay i kiu theo thi gian .................................................................... 106
5.5 Cu hi v tho lun ................................................................................ 108
Chng 6: Sinh vt tri ni
6.1 Phn loi......................................................................................................109
6.2 Cc phng php thu thp mu................................................................ 121
6.3 Cu hi v tho lun ................................................................................ 122
Chng 7: Sinh vt y
7.1 Phng php phn loi............................................................................. 123
7.2 Phng php thu thp mu....................................................................... 133
Chng 8: Dng nng lng v chu trnh khong cht
8.1 Mng li thc n.................................................................................... 136
8.2 Cc chu trnh khong cht (C, N, P)......................................................... 142
8.3 Cu hi v tho lun ................................................................................ 150
Chng 9: Cc qun x sinh vt y
9.1 Mi trng vng triu..................................................................................151
9.2 Cc vng ca sng (Estuaries) ................................................................. 154
9.3 Cc rn san h (Coral reefs)..................................................................... 159
9.4. m ly ngp mn (Mangrove swamps) ................................................ 169
9.5 bin su (Deep sea).................................................................................. 173
9.6 Cu hi v tho lun ................................................................................ 181
Chng 10: Tc ng ca con ngi
10.1. Tc ng ca con ngi n cc h sinh thi nc................................ 182
10.2. Tc ng ca con ngi n vng ca sng v vng ven bin ............... 183
10.3. nh hng ca cc hot ng con ngi n h sinh thi bin............... 187
10.4 Bo tn bin v h sinh thi bin............................................................ 193
10.5 cc h sinh thi v mi trng ven bin in hnh vit nam................. 195
10.6 Cu hi tho lun................................................................................... 200
phn 3 thc tp v bi tp ln
1. Xc nh vn nghin cu
2. Thc tp v thu thp thng tin s liu
3. ng dng
4. Lp bo co
Ti liu tham kho
Cc ph lc

Bi ging Sinh thi bin v ven b

Trng i hc Thy li H Ni- Vin nghin cu thy lc Deltf

Li gii thiu

Ti liu bi ging sinh thi bin v ven b c son tho ny l mt phn ca d
n "Nng cao nng lc o to ngnh K thut b bin ti Trng i hc Thu li".
Sinh thi bin v ven b l mt trong nhng mn hc mi trong chng trnh o to
ca ngnh ny v n s c gii thiu cho sinh vin ngnh Qun l vng b bin ca
Khoa K thut b bin thuc Trng i hc Thu li.
Mc tiu ca mn hc ny l cung cp cho sinh vin ngnh Qun l vng b bin:
(i)- S hiu bit kin thc c bn v sinh thi hc,
(ii)- Kin thc v chc nng sinh thi ca cc loi, qun th, s tng tc gia chng
v chc nng ca mng li thc n,
(iii)- S hiu bit v mi quan h gia h sinh vt v cc iu kin mi trng,
(iv)- S hiu bit v qu trnh sinh thi hc lin quan n vng b bin,
(v)- Hiu bit tm quan trng sinh thi hc ca vng b bin v m ly,
(vi)- Hiu bit p dng nhng kin thc v khi nim sinh thi hc b bin vo
thc t.
Mn hc ny c chia lm 3 phn. Phn mt gii quyt khi nim c bn v sinh
thi hc. Sinh vin s c gii thiu khi nim v cc loi, ng lc hc qun th v
chc nng qun x. Phn th hai tp trung c bit vo sinh thi bin v b bin t mi
trng sinh hc, thc vt ph du, ng vt ni, mng li thc n cho n cng ng
cc sinh vt y. Phn th ba dnh cho nghin cu tham quan thc t ng dng Vit
Nam c lin quan n nhng kin thc phn mt v phn hai.

1


PHN I

Khi qut chung v sinh thi

2


Chng 1
tng quan v Sinh thi hc
1.1 Gii thiu
Sinh thi hc xut pht t hai t ting Hy Lp oikos c ngha l nh, ni m
chng ta sng v t logos ngha l s hiu bit.
Ni chung, chng ta c th nh ngha sinh thi hc l mn khoa hc nghin cu
s nh hng ca cc sinh vt vi nhau v vi mi trng. Trong nh ngha ny mi
trng bao gm tt c cc thnh phn t nhin nh nc, t, kh hu, cc sinh vt
sng khc v cc nh hng ca chng vi nhau.
Thng thng, sinh thi hc c th chia thnh sinh thi hc trn cn v sinh thi
hc bin, trong :
+ Sinh thi hc trn cn nghin cu s nh hng ln nhau gia cc sinh vt v
mi trng trn cn ca chng.
+ Sinh thi hc bin nghin cu s nh hng ln nhau gia cc sinh vt v mi
trng t nhin ca bin v ven bin.

Hnh 1.1: S m t mi quan h qua li gia mt s yu t ca mi trng n mt
sinh vt. Trong s ny, sinh vt trung tm l ng vt n tht, nhng tng qut
ho, t con mi c hiu l thc n cho ng vt n tht cn i vi thc vt, con
mi c ngha l nh sng v cht dinh dng. S trn c n gin ho cc mi
quan h hai chiu nhng trong thc t, cc mi quan h sinh thi gia sinh vt v mi
trng l quan h a chiu.

3

Ngun gc ca sinh thi hc bt u t lch s t nhin, lu i nh tui ca loi
ngi. Nhng b tc nguyn thu ph thuc vo sn bn, nh bt c v thu lm thc
n, h cn c nhng hiu bit chi tit ni nhng con mi ca h sng u v xut
hin vo lc no. S hnh thnh nn nng nghip lm tng nhu cu cn thit nghin
cu sinh thi hc v thc vt v sinh vt nui trong nh.
Mt nh ngha chung v Sinh thi hc c nh khoa hc ngi c tn l
Ernst Haeckel ln u tin s dng vo nm 1869. ng m t sinh thi hc nh
kha cnh bn trong ca cuc sng hu c v l s hiu bit v tng mi quan h
ca cc sinh vt vi th gii vt cht bn ngoi n iu kin tn ti v c v hu c.
Charles Elton vit trong cun sch ca mnh vo nm 1972 rng sinh thi hc l
s nghin cu phn ng ca ng vt v thc vt n mi trng sng v thi quen ca
chng.
Nm 1985, Krebs nh ngha sinh thi hc l s nghin cu mt cch khoa
hc mi tng tc ln nhau quyt nh n s phn b v a dng ca cc sinh vt.
Ngy nay, sinh thi hc c hiu nh mt tr chi lp hnh khng l. y, mi
mt sinh vt ny u c nhng nhu cu i hi cho s sng m rt nhiu cc c th
khc trong vng phi hp vi chng. T , hu ht s nghin cu sinh thi hc l tr
li cho nhng cu hi V sao sinh vt ny sng hoc pht trin y m khng phi
ch khc? Hoc "V sao sinh vt hoc cc loi chng ta ang nghin cu li sng?,
Mi trng nh hng nh th no n cc sinh vt ? ngc li cc sinh vt nh hng
nh th no ti mi trng ca chng?
Tri t ca chng ta bao gm kh quyn (khng kh), thch quyn (t), thu
quyn (nc) v sinh quyn (s sng). Sinh quyn l tng hp cc vt th sng lin kt
vi mi trng ca chng.
Cng c th coi sinh quyn l ton b h sinh thi trn tri t nh v tri t, n-
c v khng kh m trong sinh vt tn ti, l tng ca cc vt th sng trn tri t.
Sinh vt hc gii quyt cc vn ca nhng thc th sng cc mc tng hp
khc nhau, t nghin cu nhng phn t sinh vt cho n nghin cu sinh quyn phc
tp. Cn sinh thi hc gii quyt c bn mi thng nht mc cao hn nh nghin cu
qun x, qun th v h sinh thi.

4


Sinh quyn
*H sinh thi
*Qun x
*Qun th
H thng sinh vt Mc tng hp tng
Cc c quan
Cc m
Cc t bo
Cc ht c quan di t bo
Phn t

1.2.1 nh ngha
Loi l mt nhm gm mt hay nhiu qun th sinh vt sinh sng m khc bit
v bn cht vi tt c sinh vt khc. Cc c th ca cng mt loi c kh nng sinh sn
ra mt th h mi kho mnh.
hoc: Loi l mt nhm cc c th ging nhau, c xu hng giao phi v sinh sn ra
th h mi kho mnh. Chng ta thng thy loi c m t khng phi bng s khc
nhau v kh nng sinh sn (mt loi sinh hc) m bi dng ca chng (thuc v mt
cu trc gii phu).
phong ph loi hay a dng loi l mt php o s loi c trong mt qun x.

1.2.2 Cc thut ng
Sinh quyn bao gm rt nhiu cc loi lin quan nhiu hoc t ln nhau. S phn
loi l khoa hc phn nhm cc loi khc nhau theo mt h thng tng tc m trong
cc mi quan h gia cc loi c quan tm nhiu nht. Cp phn loi cao nht l
cp gii. Sinh quyn bao gm nm gii.
1. Gii sinh vt ( Monera): l mt gii duy nht bao gm cc sinh vt nhn
s, chng c mt vch t bo v thiu c mng nhn ln dng a t bo. Cc
nhm khc thuc gii Monera bao gm vi khun lam (sinh vt t dng) v vi
khun tht (sinh vt d dng).
2. Gii sinh vt nguyn sinh (Protista): l gii sinh vt nhn chun lu i
nht, bao gm cc loi nhm sinh vt nhn chun (n bo thuc - a t
bo?), d dng dinh dng, t dng dinh dng v c hai dng. C l l nh
ngha chnh xc nht v sinh vt nhn chun khng phi l nm, ng vt hoc
thc vt.
5

3. Gii Nm (Fungi): L mt sinh vt nhn chun, d dng, thng l
nhm a t bo c cc t bo, c cu to a ht nhn bao gm cc t bo vi
thnh t bo. Chng ly nng lng t s phn hu xc cht, sinh vt thi ra
v hp th cht dinh dng t cc sinh vt . Mt vi nm l nguyn nhn ca
bnh tt (bnh lan nhim men, bnh g st v bnh than), trong khi nhng loi
nm khc li c ch cho vic nng, hay pha ch nh l thc n, dc phm
v cc ngun to ra thuc khng sinh.
4. Thc vt (Plantae): Thc vt khng c kh nng chuyn ng t do.
Chng l nhng sinh vt nhn chun a t bo m sn sinh ra nng lng bi
qu trnh quang hp v c cc t bo xenlulo. Thc vt l ngun sn sinh oxy,
thc n v qun o/vt liu xy dng cng nh cc loi gia v, thuc nhum v
dc phm.
5. ng vt ( Animals) l sinh vt nhn chun d dng a t bo c kh
nng chuyn ng trong sut thi gian sng ca chng, chng c nhng t bo
thiu vch ngn. ng vt cho chng ta thc n, qun o, cht bo, du thm,
c tnh gn gi thn thin v sc lao ng.
Loi l mt n v phn loi. Tt c cc c th ca mt loi u c cng
mt tn khoa hc chung. Ni chung tn ny bao gm hai t Latin. Thut ng
u tin ch h v t th hai ch tn loi. V d, Mytilus edulis (con trai xanh):
Mytilus l h ca loi ny.

6


a- Thc vt

Tp hp ngnh
(Thc vt)
275.000 loi
Lp
Angiopermae
(Thc vt c hoa)
235.000 loi

Ngnh
Tracheophyta
(Thc vt c mch)
235.000 loi

B
Rosales
Hoa hng v cc cy tng t
18.000 loi
H
Rosaceae
3.500 loi

Chi
Rosa
500 loi
Loi
Rosa gallica
Hoa hng ru

(More specific: C nhiu nt ring bit; Less specific: t nt ring bit )
Hnh 1.2: S phn loi ca thc vt. Hy ch n s lng thc vt trong mi cp
phn loi. Tp hp loi ngnh c nhiu loi thc vt hn so vi loi. Hnh nh ny
c ly t cun Cuc sng: Sinh vt hc ca Purves v nnk, ti bn ln th 4 do
Sinauer Associates v WH Freeman.

7


b- ng vt

Tp hp ngnh
(Thc vt)
275.000 loi
Lp
Angiopermae
(Thc vt c hoa)
235.000 loi

Ngnh
Tracheophyta
(Thc vt c mch)
235.000 loi

B
Rosales
Hoa hng v cc cy tng t
18.000 loi
H
Rosaceae
3.500 loi

Chi
Rosa
500 loi
Loi
Rosa gallica
Hoa hng ru
(More specific: C nhiu nt ring bit; Less specific: t nt ring bit )
Hnh 1.3 Phn loi mt loi ng vt. Hy ch n nhng im ging nhau gia tp
hp ngnh ng vt v tp hp ngnh ng vt trn. Hnh nh ny c ly t cun
Cuc sng: Sinh vt hc ca Purves v nnk, ti bn ln th 4 do Sinauer Associates
v WH Freeman.
8


C myxin
C r
Phi
Hm
Lng v
Kh Chut
Chim
b cu
Thn ln
K ging
Lng mao
v v
Vut hoc
mng
Hnh 1-4: S tin ho ca ng vt c xng sng.
1.3 Cc nhn t mi trng
1.3.1 Mi trng l g?
Ni chung, mi trng c nh ngha bi s kt hp gia cc iu kin hin ti
bao quanh nh hng n cuc sng v chc nng ca mt c th hoc cng ng. Mi
trng bao gm mi trng vt l (mi trng v sinh) v mi trng hu sinh.
Mi trng vt l biu th cc iu kin t nhin nh a cht, th nhng, a
hnh, kh hu, thi tit v thm ha, ri ro. Nhng iu kin ny nh hng ti mi
trng hu sinh v nh hng qua li vi nhau. V d nh kh hu nh hng ti loi
t.
Mi trng hu sinh l cc thnh phn sng ca mi trng (cc sinh vt khc
nhau trong h sinh thi bao gm cc c th ca cng mt loi hoc khc loi).
Bn cht ca mi trng v cc diu kin ca n rt cn thit cho s sng. Thm
ch mt ht ct trn b mt ca mt trng cng c mi trng ring. N xc nh bc x
nhn c, nhit , m hoc gi y ht ct bay i.
Hnh tinh Tri t chu bc x mt tri ti b mt tri t ph thuc vo khong
cch v chuyn ng ca n xung quanh mt tri. c trng c bit ca Tri t l do
nhit thay i trn b mt, nc c th thay i t th rn, lng sang th kh. Nc
th lng trn bn mt Tri t nh trng lc m chy t ni t cao n ni t trng.
9

S thay i trong mi trng ca Tri t ch yu c xc nh bi s bin i nng
lng nhn c t Mt tri nhng vng khc nhau v bi dng hnh hc v bn cht
ca s hnh thnh a cht.
Bc x mt tri cung cp nng lng cho tt c cc hot ng sinh hc trn tri
t. Cy xanh c th tp trung nng lng mt tri trong sinh khi nh qu trnh quang
hp. (Hnh v 1-5)

Hnh 1.5 : H c quan chnh ca thc vt
Qua , cc hot ng sinh hc trn Tri t b hn ch bi hiu xut ca qu
trnh quang hp v bc x ti. Nhng cng ca bc x cng quyt nh trng thi
ca nc, c ngha l khi no v u qu trnh quang hp xy ra, s phn b bc x
mt tri quyt nh nc c sn dng th lng. Tt c nhng iu cp trn l
nhng iu kin mi trng cn thit cho s sng trn tri t.

1.3.2 a cht v t ai
a cht vng bt k c to nn bi cc kiu dng khc nhau, y l mt
qu trnh rt lu di v phc tp, c thc hin bi s chuyn ng ca ton lc a
c gi l kin to a tng cng vi hot ng nh ni la.
a cht ca mt vng c nh hng n a hnh ca t. Cc thnh phn v c
ca t c ngun gc t b phong ho. Cc kiu c sn trong mt vng c th to
ra t bng qu trnh phong ho. b v ra thnh nhiu mnh nh do qu trnh phong
ho ho hc. Thng qua qu trnh ny, cacbon dioxyt ho tan vo nc ma to ra mt
axit nh. Loi axit ny phn ng vi b mt . Nhng qung khc nhau nh hydro
cacbon st v Magie,... l sn phm phong ho cc loi . Thng thng sn phm
R
Cung
nh sng
L
Thn cy
R
10

phong ho ca rt gn v tr to thnh chng, nhng nhiu trng hp sn phm
phong ho b xi mn v dch chuyn do dng chy mt hoc gi a ti ni khc.
vng nhit i, do vo ma kh nc mt bc hi, nc ngm ca tng su hn c
r cy trng rt ln v din bin ca sc cha nc, nn c s tp trung khong vt v
chng thm xung vo ma ma. Kt qu ca ch phong ho ny to ra mt vt
khong mu hi c gi l ong rt giu dinh dng v ngho mn. S phn b
ca cc loi thc vt thng ging nhau mt cch rt r nt vi s phn b ca cc
loi chnh.
Tt c cc loi t c to thnh t cc thnh phn chnh ging nhau. Chng l
nhng ht khong c kch thc, cu to ho hc khc nhau, v cht hu c cc giai
on phn hu khc nhau. Kt cu khong vt ca t ph thuc vo loi b phong
ho v b xi mn nh ct, bn hoc st. Tt c cc loi t l s trn ln ca cc ht c
kch thc khc nhau sp xp tng dn t cui v ht ln ri n ph sa nh hn n
cc ht st nh. Bng 1-1 ch ra h thng phn loi t ton cu ca UNESCO trong
mi quan h vi cc loi thc vt.
Bng 1: H thng phn loi t ton cu ca UNESCO
Nhm t chnh Loi t Kiu thc vt
ong (latosol) t phong ha mu Rng nhit i m
Margalite (t nhit i en) Base rich black Seasonal tropical forest
Desert soils Hot dry Desert
Chernozem Dry organic black (with
calcite)
Grasslands (steppe)
Chestnut soils Dry organic black or red-
brown
Grasslands (prairie)
Grey-brown podzol (brown
earth)
Grey-brown mull Deciduous forest
Pozol Grey banded mor Boreal conifer forest
Peats Organic peat Bog
Tundra soil or pozol (if well
drained)
Organic on mineral base
(with permafrost)
Tundra
Kiu t, kh hu v thc vt c mi lin h vi nhau. C vi h thng phn loi
c s dng rng khp th gii. qui m ton cu, kh hu dng nh yu t quyt
nh cu trc t. c im quan trng nht ca kh hu l s thay i lng ma v
nhit .
Vit Nam c khong 333.000 km
2
din tch t nhin, trong c khong 70% din
tch i ni, din tch bng phng Vit Nam phn b ch yu ng bng Sng
Hng vng min bc v ng bng Sng Cu Long min nam. (Din tch t t
nhin ca Vit Nam c khong 33 triu ha, trong c 22 triu ha t pht trin ti ch
v 11 triu ha t bi t). T l t c s dng nh sau:
- t nng nghip: 7 triu ha (chim 21%)
11

- t lm nghip: 11,8 triu ha (chim 35,7%)
- t chuyn dng: 1,4 triu ha (chim 4,2 %)
- t trng, i trc: 13 triu ha (chim 39,1 %)
Bn cnh kiu dng a cht v t nu trn, a hnh cn ng vai tr quan
trng trong vic phn b cc sinh vt (ngay c nhng thay i rt nh v a hnh).
Nhit gim theo cao a hnh c nh hng n s phn b ca cc loi sinh vt.
nhng loi t st nng trong rng, nhng ch thp trng trong nn rng tr thnh
ngp ng sau ma ln v i khi lm nc ng li, trong khi ba v nh cc dy
rng th li tng i kh. iu ny dn n s phn b ca sinh vt.

1.3.3. Kh hu (nhit , nh sng, m)
Nhit l nhn t mi trng rt quan trng. N nh hng n nhiu nhn t
khc ca mt trng v sinh v hu sinh, c bit nh nh sng, m. Mt tri l
ngun nh sng v nhit cho tri t. Khi nng lng i qua bu kh quyn, khong 20-
40% nng lng b hp th bi cc kh, bi v hi nc trong kh quyn.
S thay i ca nh sng v nhit theo cao rt phc tp. xch o, di
ngy, m hu nh bng nhau trong c nm, cn v cao hay bc cc, nam cc th
iu kin nh sng v nhit tng ma rt khc bit v khc nghit. Trong ma h
mt tri khng bao gi ln v tng khng gian chiu sng ban ngy trn 24h, cao hn
rt nhiu so vi vng nhit i. Tuy nhin, trong nhng thng ma ng mt tri rt
kh mc cao hn ng chn tri v tng thi gian chiu sng rt thp.

Hnh1-5: a) S bin i ca nhit trong nhng chu k bng gi hn 400.000 nm
trc y. b) M t s lc s pht trin trong thi k bng gi ti vng t h Rogers.
12

Chu k nhit rt quan trng vi cc sinh vt ng ng hay h. Trong s dao
ng chu k nhit , mt s sinh vt c bit s b tc ng nhiu nht bi s khc
nghit ca nng gt hoc lnh gi. C hai iu ny u nh hng n ngun nc:
Trong thi gian lnh v nc b ng bng v khng c kh nng s dng; cn trong
thi gian nng gy hn hoc bc thot hi qu mc. Nhn t khc c th nh hng
n cy trng l thi k sinh trng, cy dng nh khng c kh nng sinh trng
nhit di 0
0
C v sinh trng chm chp nhit trn im ng lnh. m v
ma cng l nhng bin s quan trng cu kh hu i vi sinh vt.

1.3.4 Cht dinh dng
Cy trng cn cung cp nng lng sinh trng v ti sinh, chng cng cn thu nhn
nhng nguyn t cu thnh nn chng. S khc nhau c bn gia s chuyn ho nng
lng v chuyn ho dinh dng l m hnh chuyn ho dinh dng v c bn l vng
trn hoc l c chu k cn s chuyn ho nng lng v c bn l khng nh hng.
Cc nguyn t cu thnh phn t m sinh vt c to thnh khng th thay i trong
iu kin t nhin trn tri t cho nn chng vn gi nguyn tnh cht khi cc phn t
chuyn t mc dinh dng ny sang mc dinh dng khc. Chng c th quay vng v
lp i lp li. Cc chu trnh cacbon, nit v photpho rt quan trng trong chu trnh dinh
dng.
Chu trnh cacbon khng phi lc no cng nm trong chu trnh dinh dng bi v
n khng cn thit phi c cc sinh vt phn hy. l do sinh vt quang hp t dng
thu nhn cacbon t kh quyn di dng cacbon ixit, trong khi cc sinh vt thi
cacbon trong qu trnh h hp.
Mc d Nit rt quan trng i vi c th sng nhng vn khng th nh lng
chi tit v chu k Nit. Cn ch rng s di chuyn ca N trong chu k nm b thiu ht
do N nm li trong bu kh quyn v trong . Trong kh quyn cha khong 4.10
21
g v
khong 2.10
23
g.
Chu trnh photpho khng tun theo mt chu trnh nht nh no. Cy trng thu
nhn photpho t t di dng H
2
PO
4
-
(pH < 7) hoc chm hn nh (HP
4
-2
) (pH > 7).
ng vt thu nhn photpho t cy trng, nu chng l loi n c, hoc t nhng ng
vt khc nu chng l loi n tht. Xc phn hu tr li photpho cho t dng photpho
st. Photpho rt kham him trong hu ht cc loi t v nc. V vy, sinh vt chc
chn mt nhiu thi gian nhn c nguyn t photpho.
Dinh dng ca cy trng:
Khng ging nh ng vt (loi sinh vt thu nhn thc n t nhng th m n n
c), cy trng thu nhn dinh dng t t v mi trng. Cy trng c kh nng to
ra tt c cc i phn t cht hu c cn thit bng cch bin i ng m chng to
thnh trong qu trnh quang hp c s dng nh sng mt tri nh ngun nng lng.
Tuy nhin, cy trng ht cc khong cht thng qua h thng r tiu th.
13

Ch dinh dng cn bng
Cacbon, hyr v xy c ht t t v l nhng cht dinh dng a lng s
cp. Canxy, magi v lu hunh l dinh dng a lng th cp cn t hn. Dinh dng
vi lng, cn rt t v i khi c hi khi lng ln. Chng bao gm: st, mangan,
ng, ch, bo v clo. Phn bn hon ho c th cung cp c ba loi dinh dng s cp,
th cp v vi lng. Nhn hiu ca phn bn s lit k cc s, chng hn 5-10-5 th
hin t l % trng lng ca dinh dng a lng s cp.
Vai tr ca t
t b phong ho, v mnh vn khong cht b phn hu ho ln vo khng
kh v nc. t mu m cha cc loi dinh dng dng d s dng m cy trng
cn phi c sinh trng. R cy hot ng nh nhng cng nhn m di chuyn trong
t v mang nhng khong cht cn thit vo b r. Cy trng tiu th cc khong cht
ny vo:
- Cc thnh phn cu to thnh cacbon hydrit v prtit
- Cc phn t cht hu c s dng trong qu trnh trao i cht, chng hn nh
magie trong cht dip lc.
- Cc hot ho Enzim nh kali, m hot ho c th so enzim.
- Duy tr cn bng thm thu.
Mycorrhizae, vi khun v khong cht
Mycorrhizae hnh thnhkhi nm (nm m hoc zygomyefe) bm xung quanh
hoc vo trong r cy v to thnh mi quan h cng sinh. Fungal hyphac ht khong
cht t t v thng qua chng ln r cy trng cn nm thu nhn hydrat cacbon t cy
trng.
Cy trng cn nit cho nhiu phn t sinh hc quan trng bao gm ht nhn v
protein. Mc d khong 70% bu kh quyn cha dinitrogen (N
2
), cy trng vn khng
th s dng trc tip c. Tuy nhin, mt vi vi khun c bit (c trong mt vi loi
thc vt) c th nhn dinitrogen (N
2
)

t bu kh quyn v bin i n sang dng nit m
cy trng c th tiu th v hp th c. Nhiu cy trng c mi quan h cng sinh
vi vi khun sng trong r ca chng: Nit hu c nh thu khong trng sinh sng.
Nhng cy trng ny c khuynh hng c cc mt r (Called Mycorrhizae) vi
khun c nh nit sinh sng.
Mi trng hu sinh
Mi trng v biu hin gen
Loi hin tng sinh hc lun b nh hng bi iu kin mi trng ca chng.
Mo Thi Lan mu ti hn cc chi, do nhit lm nh hng n s biu hin ca
loi gen. S biu hin ca loi sinh hc l kt qu tng tc gia gen v mi trng.
Mo Thi Lan v th Himalaya, c hai loi ng vt ny u c vin mu m cc
14

chi. y l do kh nng iu chnh sc t ch c th thc hin c nhit thong
hn ca cc chi . Mi trng quyt nh kiu biu hin loi sinh hc.

Hnh 1.7 : Tun lc Caribu, mt loi ng vt sng lnh nguyn

Hnh 1.8 : Rng ma n i Oa-sing-tn. Ch n mt ca cy.
1.4 Ni sng
1.4.1. Sinh thi hc c th
Sinh thi hc c th nghin cu v sinh thi, tp trung vo cc loi ring bit.
Hin nay c tnh c khong 3 n 4 triu cc loi tn ti trn mt t. Ch c rt t
trong s chng c nghin cu y chng ta hiu k v sinh thi hc c th
ca chng.
Sinh thi hc c th ca cy dng x diu hu l mt loi in hnh v n l mt
trong nhng thc vt thnh cng nht trn th gii. N mc trn hu ht cc lc a
ngoi tr Antarctica. Dng x diu hu l mt loi thch hp cho nghin cu sinh thi
hc c th. Hu ht cy dng x diu hu c cht sinh ung th, n c th l nguyn
nhn gy ung th ng vt khi n phi chng. Chng thng xuyn b hn ch bi
m, vng rm mt v khng lan rng ra xa.
15

1.4.2 Quan h gia cc iu kin mi trung v s phn b cc loi
Cc iu kin mi trng (nh nhit , nh sng, m, t, nc v a hnh)
rt quan trng i vi cc loi v n s phn b ca chng. Mi mt loi c phn ng
c th ti cc iu kin mi trng v sinh vi gi tr ti u v sc chu ng ring.
Ni chung mt loi s sng trong vng m nhng iu kin mi trng l ti u (hoc
di im cc thun) i vi n.

mn
Hnh 1-9: Phn ng ca cc loi ng vt y khc nhau vi mn

Hnh 1.10 : Tng hp tng tc gia mt sinh vt v cc thnh vt v sinh v hu sinh
ca mi trng xung quanh.
16

1.4.3 T sinh thi
T sinh thi l vai tr ca mt sinh vt chim gi v chc nng hot ng ca n
trong h sinh thi (c lin quan n cch kim n ca mt sinh vt).
Trong sinh thi, t sinh thi i din cho cch m mt loi thch nghi vi cc
iu kin mi trng ca chng. Khi nim t sinh thi rt quan trng trong sinh thi
hc v phi mt nhiu cng sc xc nh t sinh thi ca mt sinh vt mc chi
tit. Mi mt loi c t sinh thi duy nht ca ring mnh. N l mt trong s t cc
lut sinh thi. Gi s c hai loi chim gi cng mt t sinh thi song chng khng
th chim gi chnh xc ti cng mt t v cng chung sng (cng tn ti).
Ni chung, cc t sinh thi l nhng ng vt c kh nng thay i, v d nh
c th thay i khu phn n ca chng nu khu phn n a thch ca chng khng c
sn. Cc loi c chung mt t sinh thi c th tn ti chung cnh nhau.

1.4.4 Mi trng sng
Rt kh a ra mt khi nim chnh xc v thut ng ni sng, t "ni sng"
c s dng rng ri trong sinh thi m t vng m sinh vt sng. V d nh kh
t vng t thp gorilla c ni sng ca n l rng th cp nhit i. Mt s loi c
th c mt vi ni sng. Thut ng "vi cnh" (microhabitat) dng ch mt vng hn ch
ni m nhng sinh vt nh sng. Bt k mt mi trng no cng u bao gm hng
nghn vi cnh.
Theo sch Vit Nam, tng s cc loi nc ta l 10.835 loi, trong
vng b bin l 7.752 loi. S liu h thng t nhin - Ni sng v cc loi ca Vit
Nam t nm 1994 trong bng 1-3.
Bng 1-3 Ni sng v cc loi ca Vit Nam t nm 1994
V khng
xng
sng
C B st Chim V c
v
Tng
ang nguy cp 10 6 8 14 30 68
D b tn thng 24 24 19 6 23 96
ang b e do 9 13 16 32 1 71
Him 29 29 11 31 24 124
Cha xc nh 3 3 6
Tng s cc loi
dang b nguy
him
75 75 54 83 78 365
Tng s cc loi
trong nc
7000 2500 260 800 275 10835
Tng s cc loi
vng ven bin
5500 2038 10 200 4 7752

17

1.5 S thch nghi
Phn ng ca mt loi vi cc iu kin mi trng l khng n nh. Cc sinh
vt c th (trong phm vi nht nh) thch nghi vi cc thng s mi trng sng lu
i ca chng. S thch nghi l mt trong nhng c im quan trng nht ca cc vt
th sng. N cho php cc sinh vt sng st nhng iu kin mi trng khc nhau
hoc khi c s thay i cc iu kin mi trng.
S thch nghi l khuynh hng ca mt sinh vt quen vi mi trng ca mnh,
mt trong nhng im chnh ca thuyt tin ho ca Charles Darwin do s chn lc t
nhin: cc sinh vt thch nghi vi mi trng ca chng. Nhng sinh vt thch nghi tt
nht s c c hi sng st ln hn v li gen ca chng cho th h sau.
Chn lc t nhin l qu trnh sng st khc nhau v s sinh sn cc kiu di
truyn, c th n nh, nh hng, hoc ph v. Cc c th thch nghi tt hn c kh
nng sng st n tui sinh sn nhiu hn v v vy li nhiu con ci v to nn s
phn b rng hn ti ngun gen hn nhng c th km thch nghi.
Phong to thch nghi: L s pht trin ra nhiu loi khc nhau t mt gc t tin.
iu ny xy ra khi c mt mi trng sng mi thun li cho s pht trin ca mt
qun th. Kiu tin ho phn nhnh ra nhiu loi t mt t tin chung l kt qu s
thch nghi mi hoc s tuyt chng hng lot.

1.6 cu hi v tho lun

18


19
Chng 2
Qun th v qun x

2.1 ng lc hc qun th
2.1.1 Sinh sn, nhp c, dit vong, di c
Qun th l mt nhm cc c th ca cng mt loi sng trong cng mt vng,
ti mt cng mt thi im v chia s chung mt ngun gen, hay mt nhm cc sinh
vt c kh nng giao phi trong mt khu vc a l.
Cng c th hiu: Qun th l mt nhm cc c th tng t nh nhau c xu
hng giao phi vi nhau trong cng mt khng gian a l gii hn. iu ny c th
v n gin nh mt cnh ng hoa cch bit vi cc cc cnh ng khc bi qu i
hoc cc vng a l khng c nhng loi hoa ny xut hin.
ng lc hc qun th l khoa hc nghin cu cc yu t nh hng n kh
nng sinh trng, n nh, suy gim ca nhng qun th v quan h gia nhn t ny.
Mt c th l mt hoc nhiu t bo c c trng bi s sp xp thng tin DNA
duy nht. Chng c th l n bo hoc a bo. Cc c th a bo ny biu th s
chuyn mn ho ca cc loi t bo v chia thnh cc m, cc c quan v h thng cc
c quan.
Kh nng pht trin qun th lin quan n lch s s sng
Tui i ca c th sinh vt sng nh hng ti t l tng dn s. Lch s
s sng lin quan n tui trng thnh gii tnh, tui cht, v cc yu t khc
trong thi gian sng ca c th nh hng n cc c im sinh sn. Mt s
sinh vt pht trin nhanh, sinh sn nhanh v nhiu con trong mi chu k sn sinh sn.
Cc sinh vt khc pht trin chm, sinh sn mun v ch c mt vi con trong mi mt
chu k. Hu ht cc sinh vt gia hai thi cc ny.
Cu trc tui lin quan n t l tng i ca cc c th trong mi nhm tui
ca mt qun th. Cc qun th vi nhiu c th c tui sinh sn hoc trc sinh sn th
c cu trc tui dng hnh kim t thp, v c th gia tng nhanh chng v lp tr sinh
sn.
Dn s loi ngi ang trong giai on tng trng. T khi tin ho cch y
khong 200.000 nm , s lng loi ngi ca chng ta tng ln nhanh chng v lan
rng khp c Tri t. Bt u t nm 1650, dn s tng theo cp lu tha. Cc k
thut sn bn v trng trt mi gip con ngi gia tng dn s nhanh chng. Con
ngi phi mt 1800 nm t tng dn s l 1 t, nhng ch mt 130 nm t ti 2

t, v hn 45 nm t ti 4 t.
Bt chp cc tin b cng ngh, cc yu t nh hng ti gia tng dn s cui
cng s gii hn s bng n dn s ca loi ngi. Cc yu t ny gm gii hn ca
cc ngun ti nguyn vt l v sinh hc v dn s th gii vt 6 t vo nm 1999.
Nm 1987 dn s c c tnh l 5 t.
Thi
c
Bt u canh
tc nng
nghip, trng
cy v nui
ng vt trong
nh
Thi
mi
Thi
ng
Thi
st
Trung
i
Hin
i
Trc Cng nguyn Sau Cng nguyn

Cc ngnh khoa hc c
bn v tin b y hc
Bt u cch mng
cng nghip
"im cht en"
(bnh dch hch)

Hnh 2-1 : Tng trng dn s loi ngi trong 1.000 nm qua. Ch n nh hng
ca bnh tt (ci cht en) v cc tin b k thut n s dn.
2.1.2 Sinh trng qun th
ng lc hc qun th l nghin cu cc nhn t nh hng n s sinh
trng, n nh v s suy gim qun th. Tt c cc qun th u tri qua 3 giai
on ring bit trong chy k tn ti ca chng: (i)- Sinh trng; (ii)- n nh; v
(iii)- Suy gim.
- S sinh trng qun th xut hin khi cc ngun ti nguyn c sn vt

20

qu s lng cc c th c th khai thc chng. Sinh sn nhanh v t cht
thp lm cho ln qun th tng nhanh.
- S n nh qun th c bt u khi s tng trng dn s vt qu ngun
ti nguyn c sn ca chng. n nh lun l giai on lu nht trong chu
k sng ca mt qun th.
- S suy gim l s gim s lng cc c th trong mt qun th v thm ch
c th dn n s tuyt chng qun th.
Cc nhn t nh hng n s sinh trng ca qun th
Gn nh tt c cc qun th u c xu hng tng trng theo lu tha,
chng no m ngun ti nguyn cn di do. Hu ht cc qun th c kh nng
pht trin vi t l lu tha, v s sinh sn ni chung l mt qu trnh tng ln
gp nhiu ln. Cc nhn t c bn nht nh hng n t l sinh trng qun th
l t l sinh, t l cht, t l nhp c v t l di c. S thay i ln ni ti ca
qun th chnh l t l sinh cng vi t l nhp c tr i t l cht v t l di c :
E I D B N + =
trong : N l s thay i ln qun th, B l t l sinh, D l t l cht, I l t
l nhp c v E l t l di c.

B
N
D
E I

Nu chng ta b qua nhp c, di c ( I v E) v gi s rng s t l sinh trng
v t l cht l hm tuyn tnh ca ln qun th N, th ln qun th ti thi im
t + 1 c m t bi hm s:
N N
t t +
= + +
1
1 ( )
(2-1a)
hoc
N N
t +
=
1 t
(2-1b)
trong , l t l tng trng ni ti ca qun th. Hm gii tch cho cng thc (2-1b)
l:

21

t
N
dt
dN
= (2-2)
Nghim ca phng trnh vi phn ny l:
N N e
t
t
=

0

(2-3)
If > 0, Qun th tng trng theo cp lu tha;
If = 0, ln qun th l hng s trong khong thi gian;
If < 0, Qun th gim theo cp lu tha;

Hnh 2-2: Thay i ca N theo v thi gian.
Khi ngun ti nguyn tr nn b hn ch th t l tng trng ni ti () gim.
Kt qu ny l mt ng cong tng trng logic.

Thi gian

22

Mt con mi
Mt con mi Mt con mi
Loi 1
Loi 2
Loi 3
Phn hi chc nng Phn hi s lng Tng phn hi
Hnh 2-3: Ba loi thay i chnh ca qun th th n tht theo s thay i ca s lng
con mi hin c

(a)Pht trin theo lut s m
(khng hn ch c)
(b) Pht trin theo lut
Logistic
Tc gia tng
s lng sinh vt
trong qun x
Kh nng chu ng
ca mi trng
Tc
gim
xung
im gia
tng cc i
Tc
tng ln
Thi gian Thi gian

Hnh 2-4: Hai cch sinh trng ca qun th. ng cong theo lut s m (c tn
khc l ng cong J) xy ra khi khng c yu t gii hn n quy m qun x. ng
cong theo lut Logistic biu din nh hng ca yu t gii hn n quy m qun x
(trong trng hp ny l kh nng chu ng ca mi trng)

23


T
n
h

t
r
n
g

s
n
g

s
t

(
s
n
g

s
n
g

s
t
)

% trong khong thi gian
Tui (tnh theo thng)

(a) ng cong gi thit (b) Poa annua
T
n
h

t
r
n
g

s
n
g

Chim c
(c) Mt s chim hoang di (d) Cu Dall
T
n
h

t
r
n
g

s
n
g

T
n
h

t
r
n
g

s
n
g

Chim en
chu u

Chim ht
bit ht

chu M

Tui (tnh theo nm)
Tui (tnh theo nm)

Hnh 2-5: Nhng ng cong v s lng c th. a) Ba ng cong gi thit (c
nh s I, II, III); b, c, d l ba ng cong thc t. Ch rng cc ng cong thc t
gn ging vi mt trong cc ng cong gi thuyt.
S dao ng ca cc qun th gia s tng trng v s n nh
Cc yu t lm km hm n s pht trin qun th c th l: ngun thc n,
khng gian, s tng tc phc tp vi cc yu t vt l v sinh hc khc (bao gm cc
loi khc). Sau thi k pht trin theo quy lut s m ban u, mt qun th s gp mt
yu t km hm no khin cho s pht trin dng li. Qun th ny bc vo
thi k pht trin chm hn v c th cn bng mt mc tng i n nh vi bin
dao ng no . Kh nng mang (cn gi l kh nng chu ng - carrying
capacity) l im m ti kch thc mt qun th i xung.

24


M

q
u
n

x

Thi gian

Hnh 2-6: Mi quan h gia kh nng chu ng (K) v mt qun th theo thi
gian.

Dao ng theo ma
S suy gim v tuyt chng ca mt qun th
S tuyt chng l cuc u loi ca tt c cc c th trong mt nhm. Tuyt
chng cc b l s bin mt ca tt c c th trong mt qun th. Tuyt chng loi xy
ra khi tt c cc chi ca loi v cc qun th thnh phn ca n tuyt chng. Nguyn
nhn chnh ca s suy gim v tuyt chng ny l do bin i mi trng. S thay i
mt trong cc yu t vt l ca mi trng c th gy ra s suy gim v tuyt chng;
ngoi ra, ho thch cng ch ra rng mt vi s tuyt chng c th do s di tr ca i
th no gy nn.
S suy gim t ngt loi ngi xy ra trong lch s theo chu k ng vi mt
bnh truyn nhim no . Bnh dch hch git i mt na dn s chu u trong
nhng nm t 1346 n 1350, cc bnh dch khc cho n nm 1700 git i mt
phn t dn s chu ny. Bnh u ma v cc bnh khc cng lm gim i 10% dn
s khu vc Nam v Bc M.
Tc ng ca con ngi
Dn s loi ngi khng ngng tng ln do s dng cng ngh, v chnh iu
ny ph v cc qun th t nhin. S bt n nh ca cc qun th c th dn ti
nhng kt qu:
tng trng qun th khi cc yu t km hm c b ph v;
suy gim qun th khi cc yu t km hm mi t ra.
Nng nghip v chn nui l nhng v d v s pht trin ca cc qun th sinh
vt c coi l cn thit cho con ngi. Ch ring nc Anh, ngi ta nui hn
300.000 con mo, trong khi trc , t tin ca chng l mo hoang th rt him hoi,
v ch c th tm thy mt vng nh Trung ng.

25


26
nhim
Cc cht gy nhim thng l cc cht thi vo khng kh v nc mt cch
thiu k hoch v v t chc. Nhiu h thng c nit v pht pho - l nhng cht dinh
dng cho cc loi thc vt sng di nc v trn cn. Dng nc chy ra t vng
sn xut nng nghip mang theo phn bn lm tng nng nhng cht ny, dn ti
s pht trin nhanh chng ca thc vt (ph dng). Cc qun th thc vt pht trin s
dn ti s pht trin cc loi vi khun lm gim lng oxy trong nc, khin cho c v
cc loi sinh vt khc khng sng c.
Thuc tr su v s cnh tranh
Vic loi tr mt loi cnh tranh c th lm bng n s pht trin mt i th
mi. Nhng loi thuc tr su phun trn la m thng gy ra s bng n mt loi su
th cp c kh nng khng thuc cao hn khi loi su khng thuc km hn b tiu dit.
S loi tr ng vt n tht
Ngi ta thng thy ng vt n tht nhng ni con ngi t chc sn bt v
cc nhu cu khc nhau ca con ngi nh thc phm, hng da,... Khi ng vt n tht
b gim v l do no (c th do con ngi sn bt qu mc hoc bnh dch,...) th dn
ti s ng vt l mi ca chng tng ln. Chng hn nh, loi si v bo gim dn ti
loi mi t nhin ca chng nh hu, nai tng ln r rt. Ngi ta c tnh hin nay
M c nhiu hu, nai hn so vi thi m ngi chu u mi t chn ti v mt s
loi th n tht b sn bt nhiu.
Pht sinh mt loi mi
S xut hin mt loi ngoi lai nhng vng mi thng l nhn t quan trng
nht nh hng n cc qun th t nhin. V d, hn 1500 loi cn trng v 25 h c
ngoi lai xut hin Bc M; hn 3000 loi cy cng xut hin y. Phn ln
s xut hin ngu nhin sau u nhanh chng tn li, tuy nhin, mt khi mt loi
mi xut hin c s pht trin, th n s c s bng n v tng trng. V d nh
cy Kudzu, mt loi cy c du nhp vo M t Nht Bn, chim mt din tch
rng ln vng nng thn ca nc ny sau mt thi gian ngn.

2.2 S tng tc gia cc loi
Nh chng ta bit, loi l mt hay nhiu qun th cc sinh vt lai ging hay c
kh nng lai ging khc bit v bn cht vi cc sinh vt khc. Qun th ca cc c th
c kh nng lai ging v sinh sn. Cc qun th ca cc loi khc nhau sng trong
cng mt vng c th tc ng ln nhau.



27
2.2.1 S tha hip (trung lp)
y l mt kiu quan h trong hai loi khng c tc ng tch cc cng nh
tiu cc no n nhau nhau c.
2.2.3 Tnh cnh tranh
Trong qun x lun c s cnh tranh gia cc c th ca cc loi khc nhau. S
cnh tranh ny c th ng vai tr rt quan trng trong vic quyt nh mc phong
ph, sc kho, kh nng sinh sn v s phn b ca cc loi trong mt qun th. c
bit, s cnh tranh thng xy ra khi ti nguyn mi trng (thc n, nh sng, nc,
khng gian sng, tc nhn gy th phn...) tr nn khan him. Thm ch cnh tranh
cn xy ra gia cc loi tng t hay c lin quan n nhau, hoc gia cc loi hon ton
khc nhau khi chng cn chung mt ngun mi trng no . Cnh tranh l mt trong
nhng tc ng sinh hc quan trng nht nh hng n mi trng v nhng yu t
khc.
Trong nhiu trng hp, c th mt loi no khng th tn ti hay pht trin
ngay c ni c iu kin mi trng vt l thch hp cho chng: c th l do loi
ny b ln t qu mnh bi mt loi khc, tc l b cnh tranh.
2.2.4 H sinh
H sinh l mt dng ca cng sinh trong c hai loi cng c li. iu ny
th hin mt kiu cng sinh ca hai sinh vt khc loi m trong c hai sinh vt ny
cng c li. Mt v d in hnh l a y - s cng sinh gia mt loi to v mt loi
nm, trong to cung cp thc n cho nm, cn nm cung cp nc v cht dinh
dng cho to.
2.2.5 Hi sinh
Hi sinh l mt dng quan h cng sinh, trong mt loi th c li cn loi
kia th khng chu tc ng no.
2.2.6 Amensalism
Amensalism is mt dng quan h cng sinh trong cc thnh vin ca mt
qun th c ch, km hm s pht trin c mt qun th khc m khng h b mt tc
ng no.
2.2.7 K sinh
K sinh l mt dng cng sinh trong qun th mt loi c hng li do n
bm cht dinh dng t mt loi khc; tng t nh s n tht, nhng khc ch hot
ng k sinh din ra chm hn nhiu v khng phi lc no cng git cht cy ch hay
ng vt ch. Mt dng cng sinh trong mt sinh vt c hng li nh vo n
bm sinh vt khc, v d nh virus cm k sinh ngi. Cc loi virus phi k sinh
trong t bo.

2.2.8 S n tht
n tht: L mt trong cc loi tng tc sinh hc lm hn ch s pht trin ca
qun th; xy ra khi loi sinh vt ny git v tiu th sinh vt khc.
2.2.9 Quan h gia vt n tht - con mi v bin ng qun th
Quan h gia vt n tht v con mi c th c nh hng ln n s thay i v
s lng ca c hai.

Thi gian
Khong Khong
Thi gian
S con mi (N)
S
n
g

t
h
n

t
h
t

(
P
)

Hnh 2-7: M hnh vt n tht- con mi Lotka-Volterra (a) ng ng khuynh s
lng con mi , vi s con mi (n) tng (mi tn t tri sang phi) khi mt loi n
tht thp (C thp) v gim khi mt loi n tht cao. (b) ng ng khuynh th n

28

tht, vi loi n tht tng (mi tn hng ln trn) khi mt mi cao v gim khi mt
mi thp (c) Khi kt hp cc ng ng khuynh, cc mi tn cng c th kt hp,
v cc mi tn kt hp ny quay theo hng ngc chiu kim ng h. Ni cch khc,
qun th kt hp chuyn ng theo thi gian t s con mi v s con n tht cao (gc
bn tri pha di trong biu c), cho ti s con mi cao - s con n tht thp (gc
phi pha di), ti s con mi cao - s con n tht cao , ti s con n tht cao - s con
mi thp, v tr li vi s con mi thp - s con n tht thp. Tuy nhin, ch rng, s
con mi thp nht (9 gi ) i trc s con n tht thp nht (6 gi vi chuyn ng
ngc chiu kim ng h) 1/4 vng. Nhng vng trn kt hp ca cc con mi c lin
tc khng c gii hn (d). Tuy nhin, nh (e), nhng chu trnh ny th hin s n
nh: Chng tip tc mt cch khng c gii hn nu khng b gy xo trn, nhng
mi mt xo trn to nn mt mc mi s to ra mt lot chu k n nh trung
tm mi khc nhau, xung quanh cng cc gi tr trung bnh nhng vi mt bin
khc.

Gi s rng, s con mi c biu din bi cng thc sau y:
dN
dt
r N = (2-4)
S lng con mi b n tht ph thuc vo tn sut s ln chm trn gia chng.
S ln chm trn ny l mt hm ca s vt n tht (P) v s con mi (N), v mt thng
s a m t hiu qu tn cng hay ni cch khc l mc tn cng. iu ny dn ti
phng trnh vi phn ca s con mi:

dN
dt
r N a N P = (2-5)
(Quan h Lotka-Volterra ca mi)
Thiu thc n, cc con vt n tht s gy i v cht i. Nh vy s vt n tht
c gi thit l gim theo mc s m trong trng hp thiu con mi.
dP
dt
q P = (2-6)
Trong q l t l cht. S b cht c b li bi s mi sinh ra, m s mi
sinh ra ny ph thuc vo hai yu t: (1) mc thc n c tiu th, (a x N x P), v (2)
hiu sut vt n tht chuyn thc n ny thnh th h mi ca chng. iu ny dn ti
cng thc tnh cho vt n tht nh sau:
dP
dt
f a N P q P = (2-7)
(Quan h Loktka-Volterra cho loi n tht)
Cc thng s r, a, f v q c trng cho lch s cuc sng ca loi n tht v con mi.
S bin ng ca c con mi v vt n tht l mt hm ca cc thng s c
trng lch s cuc sng v s hin din giai on u ca c con mi v vt n tht.

29


30
2.3 Bc dinh dng
Bc dinh dng c th c hiu nh l s sp xp cc sinh vt vo cc nhm
qun x. Ngha en ca dinh dng l n, do vy bc dinh dng l mc n ca mt loi.
Tt c cc sinh vt u cn nng lng sng v cc sinh vt khc nhau th
np nng lng theo cc cch khc nhau. C nhiu phng php phn loi cch m cc
c th np thc n. Mt cch thng dng l chia cc sinh vt ra lm hai loi: sinh vt t
dng (hay cn gi l sinh vt sn xut), v sinh vt d dng (cn gi l sinh vt tiu
th).
2.3.1 Cc sinh vt t dng (quang dng v ho dng)
Cc loi sinh vt t dng l cc sinh vt c th tng hp cc phn t hu c
t cc hp cht v c n gin hn. Nng lng c s dng cho qu trnh ny c
ly t ngun nh sng mt tri (quang dng) hay cc phn ng ho hc (ho dng)
Cc loi sinh vt quang dng l cc sinh vt s dng nng lng mt tri
tng hp cht dinh dng hu c lm ngun nng lng ca chng; v d vi khun
xianua, to v cc loi cy. Cc phn t v c nh CO
2
, NO
3
, PO
4
c chuyn ho
thnh cc phn t sinh hc ln hn. Nhng phn t ny l cc thnh phn c bn
ca m trong c th ca sinh vt.
Cc loi sinh vt ho dng, mt vi loi vi khun c th ly nng lng t cc
phn ng ho hc, nh s xi ho sulfit hay m-tan. Nng lng ny c vi
khun s dng duy tr s pht trin.
2.3.2 Cc sinh vt d dng
Cc loi d dng l cc sinh vt np nng lng v cht dinh dng bng cch
ph v cc phn t hu c sinh ra bi cc loi t dng trong cc loi thc n; cc sinh
vt thuc loi ny bao gm cc ng vt v nm.
2.3.3 Cc sinh vt phn hu
Cc sinh vt phn hu c mt chc nng quan trng trong vic ti to chu trnh
dinh dng. Khng c cc sinh vt phn hu ny, th cc sinh vt khc trong mt qun
x s tng ln khng ngng. V cui cng th gii s cn kit Oxit carbon, Nitrat , hay
phosphate hay nhng cht v c cn thit khc cho s sng.


Khng kh
Mt tri
Cc sinh vt phn hu B thi
nh sng mt tri
Cc sinh vt sn xut
Cc cht v c
Hp cht hu c
Cc cht v c
v nc
Hp cht hu c
B cha (t hoc nc)
Hnh 2-8: Mt qun th t tn ti n gin nht cn phi c vt sn xut v vt phn
hu. Biu ny biu din mt qun th tn ti da vo nhng sinh vt quang hp t
dng v sinh vt phn hu.
Cc sinh vt phn hu ph v cc cht thi hu c v sinh vt cht thnh cc
cht hu c mi cn thit cho cc sinh vt sn xut (sinh vt t dng). Do vy, qun
x t duy tr n gin nht ch gm sinh vt sn xut v cc sinh vt phn hu, nh m
t hnh 2-8.
2.3.4 Cc ng vt n c v n tht
Cc loi ng vt n c gn lin vi cc sinh vt d dng, chng thng l cc
ng vt n c hay n to. Trong chui thc n v li thc n, cc loi ng vt n c
c vai tr nh l mt sinh vt tiu th bc 1.
2.3.5 Cc ng vt n tp
ng vt n tp l sinh vt n c thc vt v ng vt. Cc loi ng vt n tp,
theo nh nh ngha ny, l kh ph bin. iu c ngha l cc loi ny khng
chuyn vo mt loi ring bit no m tm thc n nhiu ngun a dng. Loi ngi
hu ht l n c ng v thc vt, nm cng loi thc n kh quen thuc i vi nhiu
ngi. Khi c sn ng vt, h c th sng nh vo ngun ng vt, v nu ng vt
khan him, h li chuyn sang n cc loi thc vt.
2.3.6 Chui thc n/Li thc n
Chui thc n l mt dng biu din n gin nht v nng lng trong
mt qun x. mc c bn, nng lng c d tr sinh vt t dng (thc
vt), sau cc loi n c (sinh vt tiu th bc 1) n, v n lt chng li b
cc loi tiu th bc 2 hay bc 3 n; chui thc n l mt s n gin ho, trong

31


32
hu nh cc ng vt u khng ch n mt loi sinh vt. V d loi ng vt
n tp, chng n c thc vt c ng vt.
Li thc n l mt mng li phc tp ca cc mi quan h tng tc ln
nhau v mt thu np cht dinh dng gia cc loi trong h sinh thi t nhin; Trong
li thc n, s m t v dng nng lng l phc tp v chnh xc hn so vi trong
chui thc n.
Cc h sinh thi bao gm cc vt th sng v khng sng. Nhng vt th sng
ny, hay ni cc khc l thnh phn sinh vt ny, bao gm mi trng sng v cc
sinh thi trong c cc sinh vt tn ti. Cc thnh phn khng sng, hay cn gi l
v sinh, bao gm t, nc, cc cht dinh dng v c v cc yu t thi tit. Ni sinh
sng ca mt sinh vt gi l mi trng sng. Mt sinh thi thng c xem l vai
tr c bn ca sinh vt trong qun x, l nhn t km hm i sng v s pht trin ca
sinh vt, v cn nh hng n cch sinh vt tm kim thc n nh th no.
Cc sinh vt sn xut, mt sinh thi chnh (trong tt c cc h sinh thi) l cc
sinh vt t dng, thng l quang dng. Trong cc h sinh thi trn cn, sinh vt t
dng thng l cc loi cy xanh. Trong h sinh thi nc ngt v nc mn th cc
loi to l cc sinh vt t dng ch yu.
Sinh vt tiu th (sinh vt d dng) n thc cc sn phm l cc loi sinh vt
khc. Cc loi n c l sinh vt tiu th trc tip cc loi cy c. V loi n c ly thc
n trc tip t sinh vt sn xut, chng l sinh vt tiu th bc 1. Loi n tht n ng
vt khc (loi tiu th khc), v chng l cc sinh vt tiu th bc 2 hay bc 3. ng
vt n tp (gm c loi ngi), n c thc vt v ng vt. Sinh vt phn hu l nhng
sinh vt (ch yu l vi khun v nm) ti to cht dinh dng t cc cht hu c phn
hy. Cc sinh vt phn hu ph v cc cht thi ra, cc cht hu c khng c s sng
thnh cc cht v c. Cc sinh vt nh sng trong t rt cn thit cho vi khun v nm
phn hu l v rc, chng mang li cho t ngun dinh dng phong ph.
Thm ch ngay c khi cc qun x khc nhau v cu trc, chng vn c mt s
qu trnh ging nhau nh dng nng lng (energy flow) v chu trnh vt cht (matter
cycling). Cc dng nng lng chuyn ng qua cc quan h ly thc n. Thut ng
sinh thi (ecological niche) l ch chc nng ca mt sinh vt trong mt h sinh thi.
Li thc n, chui thc n v thp thc n l 3 cch dng biu din dng nng
lng.
Sinh vt sn xut hp th nng lng mt tri v chuyn ho thnh cc lin kt
ho hc t cc cht dinh dng v c ly ra t mi trng. Ngun nng lng t thc
n hu c ri khi chui thc n; cui cng l tt c nng lng b mt i di dng
nhit; do cn c nng lng vo lin tc . Cc nguyn t v c hu ht quay tr li
t, ni m t n li c cc sinh vt tiu th s dng v khng cn mt nng
lng u vo mi no.

Dng nng lng trong cc h sinh thi, cng nh cc nng lng khc, phi
tun theo hai nh lut nhit ng hc. nh lut th nht ni rng, nng lng khng
t nhin sinh ra, cng khng t nhin mt i, m ch thay i t dng ny sang dng
khc (th nng sang ng nng). nh lut th hai l khi nng lng chuyn t dng
ny sang dng khc, mt phn nng lng s mt i di dng nhit nng. Do ,
trong bt k mt chui thc n no, nng lng s mt i khi ta chuyn dch trong
chui. Theo kinh nghim, 90% nng lng mt i khi i t mc dinh dng ny sang
mc dinh dng khc.
Ngun nng lng c bn cho hu ht s sng chnh l mt tri. Gn y, cc
nh khoa hc pht hin ra mt ngoi l l cc qun x sinh vt xung quanh cc
ming ni la phun i dng ni chui thc n bt u bng cc vi khun tng hp
ho hc -xi ho Hydro Sunphit sinh ra do phn ng ho hc v c trong v tri
t. Trong trng hp ny, ngun nng lng l lng nhit bn trong Tri t.
Cc chui thc n cho bit cc loi sinh vt no n loi no trong mt h sinh
thi, v biu din dng nng lng trong h sinh thi. Cc h sinh thi t nhin thng
c nhiu chui thc n an xen nhau. Mi bc ca sinh vt sn xut v sinh vt tiu th
l mt mc dinh dng. Mt s sinh vt tiu th bc 1 sng nh vo cy c v to nn
cc chui thc n cy c; mt s khc th n cc cht thi ra.

33

Thc vt quang hp
ng vt n c
ng vt n
tht bc 1 ng vt n
tht bc 2
Quang hp
Tiu ho, ng ho
v pht trin
Bi tit v cht

H hp
Cc qu trnh
Khng c
nhit lng
tip tc
chuyn ho
S
n
g

v
t

p
h
n

h
u

Hnh 2-9: Dng nng lng qua mt h sinh thi. Ngun: Purves v cng s, Cuc
sng: Sinh hc, S 4, Sinauer Associates v WH Freeman.

ln ca mt qun th trong h sinh thi khng chu tc ng v b km hm
bi ngun thc n, s cnh tranh, s thanh ton ln nhau v s k sinh. Li thc n

gip chng ta xc nh c hu qu ca s xo trn: nu cc loi chim n nhng con b
cnh cng v giun t, khi chng ta dng thuc tr su git nhng con b ny th s
lm tng s cnh tranh gia cc loi chim v c th lm tng s lng giun t b n tht.
Cu trc dinh dng ca mt h sinh thi to nn thp sinh thi. y ca thp
th hin bc dinh dng ca sinh vt sn xut. nh c mc tiu th cao nht, hay
cn gi l sinh vt n tht cao nht. Cc loi thp khc cng c cng nhn trong mt
h sinh thi. Thp s lng da vo s sinh vt c trong mi mt bc dinh dng.
Thp sinh khi c tnh bng cch nhn trng lng trung bnh ca sinh vt vi s
sinh vt mi bc dinh dng. Thp nng lng minh ho khi nng lng c
mi bc dinh dng k tip nhau. Thp nng lng lun cho thy nng lng gim dn
khi ta ln cc bc dinh dng cao hn, v:
Cc sinh vt bc dinh dng k tip ch tip nhn v n mt lng thc n
nht nh.
Mt s thc n c n vo nhng khng th tiu ho c v thot ra khi
b my tiu ho nh l cht thi cha b tiu ho.
Ch c mt phn thc n c tiu ho tr thnh phn vt cht ca c th
sinh vt; phn cn li c s dng nh ngun nng lng.
Phn thit yu ca nng lng thc n c s dng to nn ATP tm thi
trong c th m sau c s dng nh protein tng hp, lipit, carbon hydrat,
co bp bp tht, truyn tn hiu qua dy thn kinh v cc chc nng khc.
Ch c khong 10% nng lng sn c mi bc dinh dng c s dng
to thnh cc m (vt cht) bc k tip. Do , c th t c mt
qun th ln hn bng n ng cc thay v n cc ng vt c nui bng
ng cc, v theo cc cc nghin cu cho thy 100 kg ng cc gip ngi ta
tng 10 kg nhng nu lng ny dng nui gia sc, th cng vi thi
gian nh vy, con ngi n s gia sc ny s ch tng ln c 1 kg.

Sinh khi
(g/m
2
)
Dng nng lng
(calo/m
2
/ngy)
ng vt sn xut
ng vt n c
ng vt n tht

Hnh 2-10: Dng nng lng v t l tng i gia cc mc dinh dng khc nhau
trong chui thc n. Ngun: Purves v cng s, Cuc sng: Sinh hc, S 4, Sinauer
Associates v WH Freeman
Cc chui thc n l s n gin ho ca cc quan h phc tp. Li thc n m

34

t dng nng lng chnh xc v thc t hn l chui thc n. Li thc n l mt
mng biu din s tng tc ln nhau gia cc loi v mt thc n.
Thp thc n cho ta mt ci nhn c th v dng nng lng trong mt h sinh
thi. Bc u tin bao gm cc sinh vt sn xut (thng l thc vt). Cc bc cao hn
l cc sinh vt tiu th. Chui thc n cng ngn, th nng lng cng c sn cho cc
sinh vt.
Con ngi chim mt vai tr hng u trong cc ng vt n tht, khong 2%
nng lng c t cc sinh vt sn xut ti c cc m ca cc ng vt n tht bc 1.
S tht thot nng lng xy ra ti ni chuyn bc dinh dng. Do , trong hu ht
cc h sinh thi t nhin, cc thp thc n khng c nhiu hn 5 bc. C t ng vt n
tht ln v khng c nhiu nng lng cho chng nh thp.

Nc
Sinh vt ph
d
C nh
C n tht
Nng DDD
Chim ln

Hnh 2-11 : Lu tch sinh hc
Thc n c c nh cc sinh vt sn xut l rt khc nhau trong tng h sinh
thi. Tng nng sut s cp (NPP) l t l m ti sinh khi ca sinh vt sn xut
c to nn. Rng v m ly vng nhit i l cc h sinh thi trn cn giu kh
nng pht trin nht, trong khi h sinh thi di nc th ch c rn san h v h sinh
thi ca sng l c kh nng pht trin nht. Tt c nhng vng c cc h sinh thi
giu nh th hin nay ang gp nguy him do cc hot ng ca con ngi. Loi ngi
can thip iu chnh li khong 40% tng nng sut s cp v trc tip hoc gin tip
s dng khong 40% ton b thp thc n trn mt t. Nng lng ny khng c sn

35


36
cho cc qun th t nhin. S lu tch sinh hc xem Hnh 2-11.
2.4 Qun x
2.4.1 nh ngha v qun x
C th nh ngha qun x theo nhiu cch khc nhau, v d: Qun x l tp hp
ca tt c cc qun th sng trong mt vng no . Cc qun x c th khc nhau v
ln v ranh gii. Trong thc t, vic nhn bit cc qun x thng kh khn.
Qun x l quan h gia cc nhm thuc cc loi khc nhau. V d nh qun x
sa mc bao gm th, ch si, rn, chim, chut v thc vt nh cy xng rng sahuaro,
ocotillo v rng cy bi creozot,... Cu trc qun x c th b xo trn v thay i do
la chy, cc hot ng ca con ngi v bng n dn s; hoc: Qun x l tt c cc
loi hoc qun th sng trn cng mt vng.
Cu trc ca qun x
Cc nh sinh thi hc tm thy nhiu qun th khng c phn b mt cch
ngu nhin trong phm vi mt qun x. Trong m hnh v qu trnh phn b theo
khng gian ca cc qun th l mt thnh tu ng k ca sinh thi hc. Hai trong
nhng m hnh quan trng nht l cu trc qun x m rng v quan h him c ca
cc loi trong phm vi mt qun x.
Cc loi trong phm vi mt qun x c phm vi a l v nh im mt nh
nhau khng? Nu c, th qun x tr nn khp kn, mt n v ring r vi cc ranh
gii r rng nh l vng chuyn tip sinh thi. Tuy nhin, mt qun x m rng c cc
qun th ca n khng c vng chuyn tip v c phn b ni nhiu, ni t mt cch
ngu nhin.
Trong mt khu rng c mt cu trc qun x m th c s thay i v
m ca t. Thc vt c s chu ng khc nhau i vi cc thay i ny v chng
xut hin ti nhng ni khc nhau. u mi trng vt l c s bin i t ngt, th
chng ta thy cc ranh gii m rng gia cc qun th. V d , vng chuyn tip b
bin l vng phn bit gia nc v t (t lin v i dng). Cu trc m gip bo
v cho qun x. Ni chung khi nim v qun x m l mang tnh linh ng v t chu
chi phi hn so vi cc khi nim a ra trc kia.
Thng thng trong mt qun x c nhng loi chim u th hn hn cc loi
khc, v do ta thng gi tn cho loi cho qun x. V d rng thng (qun x
loi thng) th mang tn ca loi chim u th.
Phn loi qun x:
C hai loi qun x c bn: Qun x trn cn v qun x di nc. Hai loi
qun x c bn ny bao gm tm chnh th nh hn gi l cc qun x sinh vt. Qun


37
x sinh vt l qun x ca min rng ln bao gm nhiu qun x c iu kin t nhin
ging nhau, s phn b ca chng c iu chnh bi kh hu.
Qun x sinh vt trn cn: i nguyn, ng c, sa mc, rng n i, rng
nhit i.
Qun x sinh vt di nc: nc bin, nc ngt.
2.4.2 a dng loi v s n nh
nh ngha: a dng loi l s lng loi sinh vt sng trong phm vi mt
vng no , hoc trn ton b Tri t.
Cc qun x c cu thnh bi cc loi thch nghi vi iu kin mi trng c
th ca qun x . a dng loi v s n nh ca n gip chng ta xc nh mt
qun x v n c vai tr quan trng trong nghin cu mi trng. a dng loi ca
vng nhit i ti l cao nht v n tnh a dng loi b gim t vng nhit i ti cc
vng khc trn tri t. a dng loi l tiu chun nh gi cc loi sinh vt khc nhau
trong mt qun x (c xem nh l s phong ph cc loi). S thay i dn a dng
loi theo v cho thy phong ph cc loi gim mt cch u n tnh t vng
xch o. Mt hecta rng ma nhit i c ti 40-100 cc loi cy, trong khi mt
hecta rng n i ch c 10-30 loi cy. S ph hu mi trng sng nhng nc nhit
i trn mt n v din tch s gy nn s tuyt chng nhiu hn cc vng v cao
hn.
nhng vng nhit i, kh nng n nh mi trng ln hn, iu cho
php nhiu loi khc nhau cng pht trin. Nhng qun x vng gn xch o thng
c lch s lu i hn v chng t b xo trn bi cc hin tng bt li nh sng bng
v s thay i kh hu, do c thi gian cc loi mi tin ho. Nhng vng thuc
xch o c ma sinh trng ca sinh vt di hn cc vng xa xch o.
Trong nhng qun x di nc, s bin i dn a dng loi din ra theo chiu
su ca ngun nc. phong ph cc loi tng theo chiu su ca nc. S bin i
dn (gradient) ny c c l nh s n nh mi trng v s tng dn cc cht dinh
dng c sn trong nc.
S n nh qun x a ra kh nng duy tr s bt bin ca cc qun x theo
thi gian. Sut nhng thp nin 1950 v 1960, s n nh v s a dng loi to nn
s n nh ca cc qun x, nhng qun x c tnh a dng loi l nhng qun x n
nh. Vic ng dng m hnh ton trong nhng nm 1970 cho thy vic tnh a dng
tng, thc t c th lm tng s ph thuc ln nhau gia cc loi v dn n mt lot
tc ng khi c mt loi ch cht b loi b.
S thay i trong cc qun x theo thi gian
Cc qun x sinh hc cng nh cc sinh vt cu thnh nn chng, c th thay
i theo thi gian. Thi gian sinh thi lin quan n nhng s kin ca qun x xy ra


38
trong nhiu thp k hoc nhiu th k. Cn thi gian a cht tp trung lin quan ti
nhng s kin ko di hng nghn nm.
S bin ng ca qun x
Con ngi lun tc ng n t nhin, nh hng c bn ca hot ng con
ngi ti qun x l lm n gin ho qun x, tc l lm gim tnh a dng cc loi
(hay a dng sinh hc). Cc hot ng trong lnh vc nng nghip l s can thip c
mc ch ca con ngi m trong chng ta to ra s c canh ca mt s loi (cy
trng) a thch nh ng, la. Hu ht cc loi trong nng nghip c bt ngun t
nhng qun x tin phong.
S can thip v ca con ngi c th lm n gin cc qun x v to ra ra
nhng qun x bt buc m chng ch c mt s loi cng nh l s qu ln v s
lng ca mt s loi. S bin ng ca qun x to iu kin cho cc loi k tip c
th pht trin v sinh sn nhanh.
2.4.3 H sinh thi v qun x
H sinh thi l mi quan h ca cc nhm sinh vt nh hn vi nhau v vi mi
trng ca chng. Cc nh khoa hc thng cp ti s quan h ln nhau gia cc
thc th sng. Theo thuyt tin ha ca Darwin, cc sinh vt thch nghi vi mi trng
ca chng, chng cng phi thch nghi vi cc sinh vt khc trong mi trng .
Chng ta c th phn tch v thy rng dng nng lng i qua h sinh thi t qu trnh
quang hp ca nhng sinh vt t dng n loi n c cho n loi n tht.
C th nh ngha ngn gn: h sinh thi l mt qun x sng trong mt vng v
mi trng t nhin ca n.
H sinh thi l cp t chc qun x cao hn cng vi mi trng t nhin ca
n. H sinh thi bao gm thnh phn sinh hc v vt l nh hng n cc qun x hay
h sinh thi . Chng ta c th nghin cu h sinh thi t quan im cu trc ca s
phn b qun th hoc t quan im chc nng ca nng lng hoc cc qu trnh khc.
H sinh thi bao gm c thnh phn sng v khng sng (hu sinh v v sinh).
Nhng thnh phn hu sinh gm ni sng v t sinh thi chim gi bi cc sinh vt.
Thnh phn v sinh gm t, nc, nh sng, cht dinh dng v c v cc yu t thi
tit. Ni c tr ca mt sinh vt, ni m n c th c tm thy, chnh l ni sng ca
n. T sinh thi thng c xem nh vai tr ca sinh vt trong qun x, cc nhn
t gii hn ca s sng sinh vt v cch thc kim thc ca chng. C rt nhiu loi h
sinh thi trn th gii. Ring Vit Nam c su loi h sinh thi ch yu vng bin
v b bin:
(1)- rn san h,
(2)- ca sng,
(3)- ng bng chu th,


39
(4)- m ph ven bin,
(5)- vnh m ph,
(6)- m ly thu triu, c v c bin.
Vng chu th sng Hng v chu th sng M Cng c ng b bin ng
lc vi bn v ph sa mn bi hoc xi vi tc 10 mt mt nm. Dc di ven bin
Min Trung (T Tha Thin Hu cho n Bnh Thun) c 12 m ph. Cc m ph
ny rt quan trng cho cc h sinh thi ti cc vng .
Sinh vt sn xut, t sinh thi chnh trong tt c cc h sinh thi l sinh vt t
dng, thng l sinh vt quang dng. Trong cc h sinh thi trn cn, sinh vt sn
xut l nhng cy xanh. Trong cc h sinh thi nc ngt v nc bin, cc sinh vt
sn xut chim u th thng l cc loi to.
Sinh vt tiu th l nhng sinh vt d dng, chng tn ti c l da vo
ngun thc n ban u do cc sinh vt t dng to ra. Cc loi n c l mt dng ca
sinh vt tiu th, chng c nui sng trc tip t cy xanh (hoc loi khc ca sinh
vt t dng). Do loi n c ly thc n trc tip t cp sinh vt sn xut, nn chng
c gi l sinh vt tiu th s cp. Cc loi n tht c nui sng bi cc loi th
(sinh vt tiu th khc) v l sinh vt tiu th cp 2 hoc cp 3. Cc ng vt n tp,
con ngi thuc loi ny, n c thc vt v ng vt. Sinh vt phn hu, ch yu l vi
khun v nm ti to li cht dinh dng t cc cht hu c thi ra. Cc sinh vt phn
hu ph v cc cht hu c cht thnh cc cht v c. Cc loi sinh vt b nh sng
trong t c vai tr quan trng trong vic gip nm v vi khun phn hu rc v to
nn mu m cho t.
Cc qun x khc nhau v cu trc vn c cc qu trnh chung, l dng nng
lng v chu k vt cht. Cc dng nng lng chuyn ng qua cc quan h ly thc
n. Thut ng t sinh thi (ecological niche) l ch chc nng ca mt sinh vt trong
mt h sinh thi. Li thc n, chui thc n v thp thc n l 3 cch dng biu
din dng nng lng.
Sinh vt sn xut hp th nng lng mt tri v chuyn ho thnh cc lin kt
ho hc t cc cht dinh dng v c ly ra t mi trng. Ngun nng lng t thc
n hu c tiu hao dn trong chui thc n; cui cng tt c nng lng b mt i di
dng nhit; do cn cung cp lin tc nng lng cho h sinh thi. Cc nguyn t v c
c bn hu ht quay tr li t, ni m t n li c cc sinh vt tiu th s dng.
2.4.4 Din th ca qun x
Din th ca qun x l s thay th lin tip cc loi bi s di c ca cc loi
mi v s tuyt chng cc b ca cc loi c i km vi s xo trn dn ti hnh thnh
cc mi trng sng mi. Cc loi xm chim ban u l qun x tin phong.
Trong sut qu trnh din th, s sinh sn gim v a dng tng. Nhng khuynh

hng ny c xu hng lm tng sinh khi (tng khi lng m sng) trong mt qun
x. Din th xy ra v trong mi giai on, qun x chun b mi trng cho giai
on k tip.
Din th s cp bt u t nhng tng tr tri v phi mt mt thi gian rt
di. S phong ho do ma v gi cng vi cc hot ng ca cc loi tin phong
nh a y v ru hnh thnh nn t. Cc loi c mc vng t su hn v to ra
bng che ch cho cc loi tin phong mc thp hn st mt t. Thng v cc loi cy
rng l pht trin, qun x t cc thnh v n nh trong mi trng.
Din th th cp xy ra khi mi trng b xo trn, nh la, hot ng a cht
hoc s can thip ca con ngi (trng trt, ph rng). Loi din th ny thng bt
u vng t trng c sn cc lp t. Khc vi din th s cp phi mt mt thi
gian di to nn t, din th th cp xy ra rt nhanh. Cy tho tin phong to
ng cho mt loi cy no (v d cy thng), v ti lt loi thng c th to
ng cho rng g cng rng l.

40

Thp
Cao
M

p
h
o
n
g

p
h
Ngay sau khi khuy trn Thi gian Sau khi khuy trn mt thi
gian
Cc qun x tin phong v i u
Hnh 2-12: Din th qun x
Cc qun x cc thnh c xu hng n nh hn cc qun x ang din th. Hu
ht s tng trng v sinh sn u cc giai on din th ban u. Cc qun x tin
phong thiu s a dng, khin cho ngun vo khan him, mt nhit v mt dinh dng.
Khi s din th tip tc, s a dng loi tng ln v cc cht dinh dng c trao i
nhiu hn. Cc qun x cc thnh to nn ngun vo y hn v duy tr s n nh
ca chnh mnh v v vy m chng cng n nh hn. Cc hot ng ca con ngi
(nh ph b qun x rng cc thnh trng trt, trong c loi tin phong, nh ng
hay la) thay th cc qun x cc thnh bng cc qun x n gin hn.
Cc qun x bao gm nhiu loi, chng tin ha, v vy mi qun x cng
phi tin ha. So snh vi cc qun x bin hn 500 triu nm trc vi cc qun x


41
hin nay cho thy cc qun x hin i bao gm rt nhiu sinh vt khc nhau, c ngha
l c s tin ha v pht trin. Cc qun x hin i cng c xu hng pht trin phc
tp hn, mc d iu ny c th l s phn nh bn cht ca cc ho thch cng nh
nhng khc bit gia cc loi ho thch v cc loi sinh vt hin nay.
2.5 cu hi v tho lun

phn 2

sinh thi bin v ven b


42
Chng 3
Mi trng bin
3.1 Gii thiu chung
3.1.1 Bin v i dng (ocean)
Bin v i dng ng mt vai tr rt quan trng i vi i sng sinh vt v
con ngi. Khi ti nguyn trong t lin ang b khai thc qu mc v c nguy c cn
kit th bin v i dng tr thnh nhng ngun ti nguyn c th p ng cc nhu
cu ca con ngi. Tri t l mt hnh tinh m bin v i dng chim ti 71% din
tch b mt v c nh hng rt ln i vi kh hu ton cu. c bit bin v i
dng chim u th pha nam bn cu vi 80% din tch, cn bc bn cu ch khong
61%. Trung bnh su ca bin v i dng khong 4000 m, v hn 80% y bin
su hn 2000 m. Ni c su ln nht ca i dng khong 11000 m l ti
Marianas Trench ca i Ty Dng. Do nh sng mt tri khng th ti su hn
1000 m nn bin di su hon ton ti v lnh, thng nhit nh hn 4
0
C.
Phn trn ca bin thng m v rt giu sinh vt sng.
3.1.2 Phn b ca bin v i dng
Bin v i dng ca th gii c th c chia thnh cc i dng v bin ven
b. Thi Bnh Dng l i dng ln nht v chu nh hng tng i t bi s thay
i kh hu gia cc vng hoc dng chy t cc sng t t lin xung quanh. Cc
qun o tp trung nhiu nht Thi Bnh Dng, cc hot ng ni la cng xy ra
nhiu nht y v vng xung quanh. V d ni la St. Helens Washington, Fuji
Nht Bn, Krakatoa ty nam Thi Bnh Dng v iu ny gip ta hiu rng ti sao
ni Thi Bnh Dng l vng cung la.
i Ty Dng tng i hp v c bao bc bi cc bin ven b (v d, vnh
Mexico, bin a Trung Hi, bin Bantic, bin Bc). i Ty Dng l ni nhn nc
ca cc con sng ln nht th gii (v d, sng Mississippi, Amazon, Nile, Congo).
su ca n nh hn Thi Bnh Dng. i Ty Dng chu nh hng vi mc ln
hn bi kh hu lc a v cc cht ho tan t cc sng vo. Ranh gii pha bc ca
i Ty dng l hi t cn nhit i, ni nc lnh v mn hn truyn v pha bc.
Ni chung gi ty trong khong 40 60
0
v nam to ra dng hi lu trn b
mt v pha ng xung quanh Nam cc. Hi lu ny v cc hi lu quan trng khc
c biu th hnh v 3-1.
Hu ht cc bin ven b c cc c trng hi dng hc ring, do chng c s
kt ni mt phn vi i dng m v su ca chng thng nh. V d, vng nc
nng gn Gibraltar hn ch s trao i gia a Trung Hi v i Ty dng v do
bc hi vt qu lng ma nn mn ca a Trung Hi tng ln. R rng trong
thi k a cht gn y (5-6 triu nm trc), s h thp mc nc bin ni lin vi


i Ty Dng v do bc hi ln rng khp to ra s lng ng mui quy m ln.
Hu ht cc bin ven b c mt lch s tng t do s thay i mnh m cc iu
kin t nhin.

Hnh 3-1 Cc dng hi lu b mt ca cc i dng (Ngun: Hi lu ca cc i
dng, Walter Munk. Copyright 1955 by Scientific America. Inc. All rightss reserved)

43


44
3.1.3 Cc dng hi lu ca i dng ( )
ng s xo trn ca cc lung gi v s khc
nhau ca mt nc cc tng nc.
1- Cc dng nc mt ca i dng (The oceanic surface currents ):
to ra s lch hng v pha tri i
vi nc chuyn ng Nam bn cu (hnh 3-3).
hiu kim ng h Bc bn cu v theo chiu
ngc kim ng h Nam bn cu.
n ny lm lch hng chuyn ng ca cc
khi nc mt v pha ty (hnh v 3-5).
c bit nh l hin tng ElNino. N phn nh mt chu trnh mang tnh
ton cu.

Ocean circulation
Cc dng nc trong cc bin h c xem xt hai kha cnh: cc dng mt
v dng thng ng (theo chiu ngang v chiu su). Cc dng mt chu tc ng bi
cc tng tc ca h thng gi ca hnh tinh v chuyn ng quay ca tri t. Dng
thng ng v dng nc su chu tc
Hot ng quay ca tri t to ra s lch hng ca dng mt c gi l hiu
ng Coriolis, n nh hng ti dng nc trong cc quy m a l khc nhau. Tri t
quay mt vng trong mt ngy xung quanh trc ca n. Mt phn t trn b mt tri
t ti xch o phi chuyn ng t ty sang ng nhanh hn mt phn t ti v tr gn
cc v n phi i qua mt khong cch di hn khong cch quay mt ngy ca tri
t. Hiu qu quay ny c gi l hiu ng Coriolis. N to ra s lch hng v pha
phi i vi nc chuyn ng Bc bn cu v
Tnh cht d thy nht ca mt i dng quy m ln l s c mt ca cc
dng trn mt, n chu tc ng bi h thng gi ca hnh tinh tri t (hnh 3-4). Gi
c to thnh do s thng cao ca khng kh nng di tc ng ca nng lng mt
tri v s ging xung ca khi khng kh khi b lnh i. Cc chuyn ng ny chu
nh hng ca hot ng quay ca tri t. Trong khong 30 60
0
v bc v nam,
gi ty chuyn khng kh v pha ng bc (Bc bn cu) hoc v pha ng nam (Nam
bn cu). Gia xch o v v 30 bc v nam, gi mu dch thi v ty nam (Bc
bn cu) hoc thi v ty bc (Nam bn cu). Cc loi gi ny kt hp vi hot ng
trc tip ca hiu ng Coriolis di chuyn nhng khi nc mt khng l vi quy m
ln nh l nhng vng xoy v i, tp trung khong 30 v bc hoc nam. Cc
vng xoy ny chuyn ng theo c
Ti cc i dng m, gi v hiu ng Coriolis kt hp vi nhau to thnh
"ging ngc" (upwelling), n mang nc di su giu dinh dng ln b mt. Hin
tng ny xut hin khi hiu ng Coriolis nh hng mnh n chuyn ng ca
nc cc thm chu lc ca pha ng cc i dng kt hp vi gi thi trc tip t
hng bc hoc nam. Nhng nguyn nh
Thng thng c vi nm th s o ngc lin tc ca cu trc dng chy xut
hin pha ng Thi Bnh Dng to nn s kt thc c bn ca hin tng "ging
ngc" v



Hnh 3-3 Hiu ng Coriolis nh hng ti cc vt th chuyn ng nh phn t nc
trong hnh v. Mi tn t nt biu th phn t chuyn ng khng b lch hng, mi
tn lin nt biu th phn t chuyn ng chu s lch hng. Trn xch o chuyn
ng khng b lch hng, cng xa xch o chuyn ng cng lch hng.
S chuyn ng mnh l thng ca nc mt c to nn bi sng do bo v
ng t. Nhiu s kin bin i hon ton khc c th gy n tng su sc nh hin
tng El Nino rng ln trn Thi Bnh Dng. Cc cn bo xoy l rt ph bin c
i Ty Dng v Thi Bnh Dng v chng c th ko di trong vi tun. Nhn tng
th nhng h thng bo c th di chuyn vi tc vi cy s mi gi v c th bao
ph mt din tch hng ngn cy s vung. Tc gi c th ln n 160 km/h v sng
dng c 10 m vng ven bin l iu c th. Nhng cn cung phong c th ph hu
hon ton mt di san h v to ra nhiu tc ng ti t khc. V d, nm 1989 trn
cung phong Hugo vi sc cng ph cc mnh ht tung nhng khi ln v
nh ca ca vng Charleston v Nam Carolina. Hay nhng trn ng t hoc lng
ng nhanh ca bn ct ph sa cng c th to ra nhng tc ng rt ln. Chng c th
to ra nhng t sng gi l sng thn, sng ny c th di chuyn vi tc trn 600
km/h. cao sng c th rt nh trong bin h, nhng nu vo vng ven b bin nng
th chng to ra nhng ct nc cao v c th to nn nhng sng nc dng ti 5 m.
Vo cui th k XIX, hng chc ngn ngi thuc cc vng b bin ca Nht Bn
b cht do mt trn sng thn c tnh ph hu c bit. Trn ng t gy sng thn
ng Nam A' u nm 2005 cun i v lm cht hng trm ngn ngui cc nc
Indonexia, Thi Lan,... Khc phc cc hu qu phi mt hng ngn t la v hng
chc nm.

45



Hnh 3-4: Mi quan h ca cc dng mt ca i dng vi h thng gi ca hnh tinh
(Ngun: Fleming, 1957, courtesy of the Geological Society of America.)

Hnh 3-5 Hin tng ging ngc ca nc ven b c to nn bi gi kt hp vi
hiu ng Coriolis.

Trong trng hp vnh hp, s lu thng trao i gia bin h v vnh c biu
th nh hnh 3-6.
46



Hnh 3-6 S lu thng gia bin m v vnh hp. Thiu s trao i ca nc y vnh,
kt hp vi h hp lm cho nc thiu xy
2- S lu thng theo chiu ng v lu thng nc bin su
Trong cc bin h, s lu thng theo chiu thng ng vng nc su c
thc hin ch yu do s xo trn ca gi v s khc bit gia mt nc cc tng
khc nhau. Ngc li vi cc c trng ca dng mt, s lu thng ca nc bin su
c c trng bi s chuyn ng ca cc khi nc ln ng nht v nhit v cc
c trng mn mt bin ti cc v cao. Cc khi nc v cao chm xung
do khi lng ring ln v chuyn ng ti cc v thp. Pha di mt nc bin,
cc khi nc c cc gi tr khc nhau c xc nh bi khi lng ring ca cc khi
nc. Cc khi nc ny c chng ln ging nh mt ci bnh xp theo lp, theo th
t gim dn ca s gia tng khi lng ring nc. Khi lng ring nc bin l
mt tnh cht quan trng nh hng ti s chuyn ng theo chiu thng ng ca
nc, n b nh hng ch yu bi mn v nhit . Khi lng ring nc bin
c biu th bng s gam trn mi cm
3
. Khi lng ring tng khi nhit gim (ln
nht 4
0
C) v mui tng. Khi lng ring nc bin ch ln hn vi phn trm so
vi nc ct, thng thay i gia 1,02 n 1,07 g/cm
3
.
S bin i mnh m khi lng ring nc bin theo chiu su xut hin cc
mc khc nhau trong i dng. Bin i theo chiu su ca khi lng ring nc
bin c bit nh l t trng (pycnoclines). Trong ma h, mt tri lm nng t
pha trn mt bin to nn mt lp nc m dy khong 3-10 m. Khi ma thu n, s
phn tng ny b ph v v gi d dng lm xo trn cc lp nc cho ti khi ma
ng lm gim nhit nc v kt qu to ra ct nc c xo trn u v cng
nhit lnh t mt n y bin.
Trong phm vi khng gian ln nht, bin nhit i bao gm mt lp nc m b
mt, mt lp trao i nhit gia v mt lp nc lnh tng su. Ti lp mt, mt
tri cung cp nhit quanh nm, lp nc su c gi lnh nh s lu thng tng su.
Cn bng gia ma v bc hi trong vng nhit i dn ti gim mt nc b mt.
Hnh 3-7 biu th quan h in hnh gia nhit v su ca i dng.

47



Hnh 3-7 : Quan h gia nhit nc bin v su.

Cu trc thng ng ca i Ty dng (Hnh 3-8) c gii thch nh sau. Ti
bin Weddell ca Nam cc, khi lng ring cao hn ca nc lnh hn c tng cao
bi mn tng ln do ng bng. V vy nc c khi lng ring rt cao y Nam
cc c hnh thnh, n chm xung v chuyn ng dc theo y bin ti bn cu
Bc. Nc su Bc i Ty dng (NADW) to nn bin Na Uy nhng khi lng
ring khng nh nc y Nam cc. N chm xung v chy theo hng nam,
chuyn ng trn nc y Nam cc. Nc gia Nam cc (AAIW) c to thnh
gn vng Nam cc nhng n khng ch lnh v mn nh nc su Bc i Ty Dng.
N chm xung, nhng di chuyn ti phn cn li ngay pha di lp mt vng nhit
i ca nc c mt thp v m, v trn lp nc su Bc i Ty Dng. Nhng
chuyn ng ny to nn mt chui cc lp nc c khi lng ring thp tng dn khi
mt lp chuyn ng ln pha mt bin. Tt nhin c s xo trn ng k ti khu vc
gia cc khi nc, nhng tuy nhin cc khi nc c th c nhn bit bi cc t
hp c trng v mn v nhit . Thi gian ca chuyn ng c th ko di hng
ngn nm.
Ngun gc ca cc khi nc ny c bit quan trng i vi s tn ti ca s
sng tng nc su bi v nc bt u t b mt bin c bo ho xy. S tiu th
xy sut trong qu trnh vn chuyn ca khi nc khng lm cho nc thiu
xy, v v vy bin su vn c xy ho ng k. iu ny cho php sinh vt di
bin su c s cung cp xy kh phong ph.

48



Hnh 3-8 S lu thng nhit nc su ca i Ty Dng. AABW- nc y i Ty
dng, AAIW- nc gia i Ty dng, NADW- nc su Bc i Ty dng
(Ngun: Gerhard Neumann and Willard J. Pierson Jr., Principles of Pbysical
Oceanograpby, copyright (D 1966. Reprinted by permission of Prentice-Hall, Inc.)

3.1.4 Mt ct ngang ca i dng
a hnh v cu trc ca y i dng: Cc i dng c 3 tnh cht v a
hnh ch yu: (i)- mi dc v thm lc a; (ii)- y bin su; v (iii)- cc h thng ni
i dng.
Thm lc a c dc nh (1:500, khong 1 ), c dng bng phng ko di
khong 10 km n 300 km t ng b (hnh 3-9). V pha bin ch gy ca thm dc
(thng su 100 200 m), dc ca lc a tng ti khong 1:20 (khong 2
0
9).
dc lc a thng b chia ct bi cc hm su vi vai tr nh l nhng knh ti bn
ct. Chn sn dc kt hp vi s nng ln ca lc a to nn s dc xung 2 - 4 km
ti phn t phng su thm vi su trung bnh khong 4000 m. Ti mt s vng
ca i dng (v d, b bin pha ty ca Nam M), nhng rnh su xut hin ngay ti
pha bin c th su ti hn 10.000 m. Nhng rnh su ny di v hp chy song song
vi b bin. Nhng o c lp gia i dng c th mc ln t y bin su ti mt
nc (v d, o Hawaii). Ngc li, cc o lc a c hnh thnh t thm lc a.
Cc i dng trn th gii c th c chia ra mt s mi trng bin khc
nhau. Phn loi c bn nht l tch bit mi trng sinh vt bin khi v sinh vt y.
Mi trng bin khi l nhng g thuc khi nc bin t b mt n su ln nht.
Cn mi trng y bao gm y bin v cc vng b bin, ven bin hoc cc vng
thy triu, cc rn san h,...

49



Hnh 3-9 Cc vng sinh thi c bn ca i dng. Vng bin ven b c tch bit
khi i dng bi g lc a khong 200 m su.

Mt cch phn chia khc tch bit i dng rng ln thnh mi trng i
dng t vng ven b. Cch chia ny da trn su v khong cch t t lin, v
vic tch bit mt cch tin li l ti gii hn su 200 m ni thng nh du g
ca thm lc a (hnh 3-9). Ti mt s vng nh b bin pha ty Nam M ni m
thm lc a rt hp, th khu vc bin ven b ch ko di mt khong rt ngn t b
bin. Ti cc ni khc (b bin ng bc nc M), th vng bin ven b c th ko di
hng trm km t t lin. Ni chung, cc thm lc a ch chim cha ti 8% din tch
cc i dng, din tch ny ch tng ng din tch chu u v Nam M cng li.
Cc sinh vt bin c th c phn loi theo mi trng bin m chng sng. V
vy c cc loi sinh vt i dng v cc loi sinh vt bin ven b tu thuc vo cc
sinh vt ny c tm thy vng i dng xa b hay vng nc ven b. Mt cch
tng t, cc ng vt v thc vt sng cng nhau y bin c gi l sinh vt y.
Sinh vt y bao gm rong to bin, cc ng vt khng xng sng nh bt bin, cc
ng vt ny b trn nn y bin hoc cc hang.
Mt ct ngang ca bin t ven b bin n bin su c th chia ra 6 phn vi cc
tnh cht khc nhau (hnh 3-9). Trong , chng ta c th thy vng ven b
(supratittotal, phn trn mc nc cao ca bin), vng st b (vng thy triu), vng
gn b (sublittoral), .... v vng bin su trn 6000 m (hnh 3-9).
3.1.5 Cc tnh cht vt l v ho hc ca nc bin
Ni chung, cc tnh cht vt l, ho hc c bn ca nc gm:
- Sc cng mt ngoi (Surface tension);
50


- Tnh dn nhit (Conduction of heat);
- Tnh nht (Viscosity);
- Nhit chuyn trng thi: lng nhit thu vo hay mt i trn mt n v khi
lng do vt cht thay i t trng thi lng sang kh hoc t kh sang lng m
khng c s tng nhit (cal/g);
- Nhit dung (Heat capacity): Lng nhit yu cu tng nhit ln 1
O
C ca 1
gam vt cht (cal/g/
O
C);
- Khi lng ring (density): Khi lng trn 1 n v th tch (g/cm
3
hoc g/ml);
- Kh nng ho tan ().

Lin kt
Hyro

Hnh 3-10 Phn t nc. Tng in tch m mt bn v tng in tch dng mt bn
khc l kh nng c bn ca nc hnh thnh dng lin quan vi ion, lm cho nc
c kh nng ho tan tt.

Nc bin l nc trong ho tan mt s cht rn v kh khc nhau. Trong
1000 gam nc bin c cha khong 35 g hp cht ho tan, c gi chung l cc
mui. Ni cch khc, 96,5% nc bin l nc v 3,5% l cc cht ho tan. Tng
lng khong ho tan c gi vi thut ng l " mn". Cc nh sinh hc bin v hi
dng hc trc y thng biu th mn di dng phn ngn, c vit tt l %o,
nhng thut nh mi hin nay dng l n v mn thc t (practical salinity units -
psu). V vy, nu mt mu nc bin in hnh c 35 g hp cht ho tan trong 1000 g,
th n c mn l 35 psu (35%o).
Cc cht ho tan bao gm cc mui v c (chim phn ln nht), cc hp cht
hu c nhn c t cc sinh vt sng v cc kh ga ho tan. Cc mui v c trong
nc bin tn ti di dng cc ion. Su ion v c chim 99,28% trng lng ca cht
rn ha tan trong nc bin. Chng l clo, natri, lu hunh (nh sulfate), magi, nhm,
v cc ion kali (xem bng 3-1). Cc cht v c ny c xem l cc ion chnh. Bn ion
ph lm tng thm khong 0,71% trng lng v nh vy c 10 ion kt hp vi nhau
to nn 99,99% trng lng ca cc cht ho tan.
51


52
mn nc bin cc khu vc khc nhau ca i dng cch xa vng b thay
i trong mt khong rt hp, thng thng l 34 37 psu, v trung bnh l 35 psu.
mn khc nhau l do s khc nhau v lng ma v bc hi. Nhng mn cao hn
xut hin cc i dng cn nhit i, ni c lng bc hi ln v mn thp cc
i dng c nhit n ho, ni m lng bc hi nh hn. mn vng xch o
thp hn v lng ma ln trong khu vc hi t gia hai ch tuyn. Ti cc vng ven b
v cc bin kn cc b, mn c kh nng thay i nhiu hn v c th t gn 0 psu
ni c cc sng ln nc ngt vo, v t ti 40 psu nh bin v Vnh Ba T.
Mc d cc thay i mn t 0 n 40 psu, cc t l trong cc ion chnh vn
khng i theo chiu thng ng. iu ny rt quan trng, c ngha l nng hu ht
cc ion trong mt mu nc c th c xc nh bng vic o c mt ln. Quan h
c bn ny l c s cho vic o c mn ca nc bin. mn c th c xc
nh bng vic o c mt c tnh ring l, c th l Clo (nng ion Clo), hoc
dn in hay ch s khc x m c tnh ca n cng ph thuc vo nng mn.
Trong phn cn li 0,01% ca cc cht ho tan trong nc bin gm mt s
mui v c m chng c tnh quyt nh ti sinh vt vin. Bao gm cc cht dinh
dng v c, pht pht v natri, chng rt cn cho thc vt tng hp cc cht hu c
trong quang hp, v dioxit silic cn cho to ct v cho cc sinh vt to xng. Ngc
vi cc ion, nitrat (NO
3
-
) v pht pht (PO
4
3-
) khng tn ti theo mt t l nht nh vi
cc nguyn t khc hoc ion khc. Chng c ngun nc mt cung cp, thay i
nng do hot ng sinh hc ca sinh vt. Trong mt s trng hp, s cung cp cc
dinh dng thit yu c th lm hn ch s pht trin ca thc vt.
Bng 3-1: Cc thnh phn chnh v ph ca nc bin 34,8 psu
Ion Phn trm trng lng
A. Cc ion hnh
Chloride (Cl
-
) 55.04
Sodium (Na
+
) 30.61
Sulfate (SO
4
2-
) 7.68
Magnesium (Mg
2+
) 3.69
Calcium (Ca
2+
) 1.16
Potassium (K
+
) 1.10
Subtotal 99.28
B. Cc ion ph
Bicarbonate (HCO
3
-
) 0.41
Bromide (Br
-
) 0.19
Boric acid (H
3
BO
3
) 0.07


53
Strontium (Sr
2+
) 0.04
Subtotal 0.71
Tng cng 99.99
Ngun: From Deep Sea Biology: Developments and Perspectives by N. B.
Marshall, C 1980 Garland STPM Press.

Cc cht cn thit khc i vi s tn ti qu trnh sng l cc cht vi lng
gm cc yu t nh st, mangan, co ban, v ng, trong quan trng l st. Theo
Matin v cng s (1994), trong mt s vng bin trn th gii ni m nitrat v pht
pht rt phong ph, sn lng thc vt vn b gii hn bi thiu cht vi lng ca st
trong nc (V d, trng hp trong nc Nam cc). Mt s hp cht hu c nh cc
vitamin cng c coi l cc cht vi lng nhng t c bit v s thay i ca chng.
Nng mui ca nc bin c nh hng nht nh n cc tnh cht ca
nc. Khi lng ring ln nht ca nc tinh khit l 4
O
C, nhng trong tt c cc
loi nc c mn trn 24 psu, khi lng ring tip tc tng ln n im bng. Do
nng mui im ng bng cng gim xung t 0
O
C, gim l hm s ca
mui. i vi nc bin 35 psu th im ng bng l -1,9
O
C. Tuy nhin trn nhit
ng bng, cc mui b tch ra v khi lng ring gim xung; v vy bng tri trn
mt nc. Tm quan trng ca gia tng khi lng ring ny l lm cho nc mt rt
lnh v khi lng ring rt cao, nc ny thng c gi l nc mui v c th
c hnh thnh v chm xung y bin, ti n hnh thnh mi trng cho cc sinh
vt bin su v quan trng hn l to nn s lu thng rt mnh lm thng kh cho bin
su.
xy v ioxit cac bon l hai cht kh ho tan trong nc bin c tm quan trng
i vi trao i cht. Kh nng ha tan ca kh trong nc bin l hm s ca nhit ;
nhit thp th kh nng tan ca kh ln. V vy, nc lnh hn c nhiu xy ho tan
hn. D ng nh vy, kh nng tan ca kh trong nc l khng ln. Ti nhit 0
O
C,
nc bin 35 psu cha khong 8 ml xy trong 1 lt nc, trong khi khng kh c th
cha 210 ml/lit. Ti nhit 20
O
C, nc bin 35 psu ch cha 5,4 ml/lit. Nguyn nhn
nc bin su khng thiu xy (devoid of oxygen) mc d cc hot ng sinh hc vn
xy ra khi nc chy qua y bin l v nc bin su c chm t nc mt, n rt
lnh v c lng xy ho tan ln nht, ln hn c lng xy b tiu th bi ng vt t
i bin su. xy phn b khng ng u theo su ca i dng. Din bin theo
su cho thy hm lng xy ln nht l trn lp nc 10 - 20 m do hot ng
quang hp ca thc vt lm cho bo ho xy. su tng ln th hm lng xy suy
gim, t ti gi tr nh nht trong khong su 200 -1000 m cc i dng m.
Ti vng ny xy c th t gi tr bng khng mt vi ni. Bn di vng ny, hm
lng xy li tng ln mt t theo chiu su nhng chng thng khng th bng gi
tr trn lp mt. S xut hin vng xy nh nht l do hot ng sinh hc lm suy
gim xy, hn na do thiu hot ng quang hp v tip xc vi kh quyn phc


54
hi xy. xy tng ln pha di vng xy nh nht l kt qu t cc ngun xm nhp
ca nc giu xy b chm xung y i dng nhng v cao.
Kh nng ho tan ca dioxit cacbon c hi khc so vi xy, v n phn ng ho
hc trong nc. Nc bin c rt nhiu dioxit cacbon, n c kh nng hp th rt tt
CO
2
. Nguyn nhn l do khi tan vo nc bin, dioxit cacbon phn ng vi nc to
ra a xit cacbonic nh sau:
CO
2
+ HOH H
2
CO
3
(3-1a)
A xit cacbonic sau phn ly thnh ion hydro v ion bicarbonat:
H
2
CO
3
H
+
+ HCO
3
(3-1b)
Bicarbonat sau c th phn ly thnh ion hydro khc v mt ion carbonat:
HCO
3
-
H
+
+ CO
3
2-
(3-1c)

Mt kho cha khng l dioxit cacbon trong i dng l ion bicacbonat, c th
thy v tr u th ca n trong cc ion ph trong nc bin (bng 3-1). Ngc li vi
oxy trong nc bin, CO
2
li chim u th hn l trong khng kh. V vy, s c sn
ca dioxit cacbon l khng hn ch i vi pht trin thc vt trong nc bin.
H thng axit- bicacbonat dioxit cacbonic l mt h ho hc phc tp dn ti s
cn bng. Nu kh CO
2
b tiu hao t nc bin th cn bng s b ph v, v axit
cacbonic s thay i v pha tri ca cc phng trnh trn, cho n khi nhiu CO
2

c to thnh v mt cn bng mi li c thit lp.
Cc phn ng cho thy kt qu to ra cc ion hydro t do (H
+
). S phong ph
ca ion hydro trong dung dch l thc o a xit. C nhiu ion H
+
c ngha l dung
dch nhiu a xit, v c t ion H
+
c ngha l dung dch c tnh kim. Cc dung dch
kim l nhng dung dch c mt s ln ion OH
-
v t ion H
+
. Tnh a xit v kim c
tnh theo h logarit t 1 n 14 n v. Nng ion H
+
cao th pH nh; cn gi tr
pH nh biu th iu kin a xit. Ngc li khi pH cao tc biu th nng H
+
thp v
OH
-
cao. im trung tnh vi pH = 7, s ion ca c hai loi bng nhau.
Mc d nc tinh khit l trung tnh v s phn ly ca phn t nc to ra s
ion H
+
v OH
-
c nng bng nhau, s c mt ca CO
2
v cc ion kim mnh nh
natri, kali v can xi trong nc bin dn ti s thay i cn bng. Nc bin c tnh
kim nh, thng thng pH khong 7,5 n 8,4. Vi pH ca nc bin, h thng
axit- bicacbonat dioxit cacbonic ng vai tr nh l mt h m gi pH ca nc
bin nm trong khong thay i nh. H axit ny s hp th H
+
khi nng ca chng
vt qu gi tr cn bng v to ra H
+
nhiu hn khi chng t c cung cp. N c
tin hnh nh thay i cc phn ng nh nu trn sang pha phi khi c qu t ion
H
+
, to ra nhiu hn cc ion bicacbonat v ion cc bonat, v chuyn phn ng sang tri
khi c qu nhiu ion H
+
, to ra nhiu hn axit cacbonic, dioxit cacbon v cc ion
bicacbonat.


55
3.2 cc c trng v sinh ca nc bin
Trong mi trng bin c nhiu rng buc vt l v ho hc, chng c nhiu
nh hng ln n cc h sinh thi ca bin. Mt s rng buc v sinh thi l do bn
cht t nhin ca nc bin vi nhng tnh cht c bit ca nc v cc ho cht ho
tan trong n. Mi trng v sinh rt quan trng i vi cc sinh vt c tr trong bin.
3.2.1 Bc x mt tri (Solar radirion )
Nng lng nh sng ti tri t bt ngun t mt tri, v v vy hu ht nh
sng ti bin t pha trn, ti b mt bin. Nng lng nh sng bin i trong khong
t nh sng ngn nh tia cc tm n nh sng quang ph c th nhn thy c, n
nh sng hng ngoi v quang ph sng di.
Bi v bc x mt tri l ngun nng lng cho quang hp, nn nh sng l yu
t quyt nh i vi s sng. Hu ht cc ng vt bin cng ph thuc vo nh sng
di chuyn, tm mi v nhn dng i th. Cng nh sng gn b mt nc
lm suy gim hot ng sinh hc thng qua vic lm mt tc dng ca protein v
DNA. Tia cc tm c hi, c bit i vi nc trong ca vng nhit i, ni m mt
lng nh sng ng k s xuyn ti su 10 m. Mt s loi thc vt ca vng thy
triu s dng canxi cacbonat gim tc hi ca tia cc tm bng cch hp th chng.
Mt s san h c sc t c kh nng hp th tia cc tm. Tia cc tm khng gy hi
trong nc c nhiu v hn 90% tia cc tm b hp th trong lp nc trn mt.
nh hng ca nh sng mt tri b gim nhanh chng theo chiu su. Trong
vng nc c ven b nh sng nh ra gii hn i vi s pht trin ca thc vt thm
ch su 10 m. Ti nhng su 30 - 50 m, kh nng nhn thy b gim i rt ln.
iu ny c bit ng i vi s phn bit mu sc, v cc ng vt sng nc su
khng c kh nng nhn thy mu sc tt. Trong bin h, ti cc su ln hn 1.000
m c rt t nh sng mt tri. Cc ng vt th gii i dng su ch thy tia sng
m nht t pha trn.
1- Thay i bc x theo ma v v
Cc nh sinh hc i dng c xu hng s dng mt s thay i cc n v
o c bc x mt tri ti mt bin v cng nh sng ti cc su trong bin.
Bc x mt tri n kh quyn bn ngoi tri t l khng i v c gi tr vo
khong 1360 watt trn m
2
(hoc 1,94 calo/cm
2
mi pht) (hnh 3-11). Khong mt na
nng lng ny b hp th v ri ra trong cc lp kh quyn khc nhau, v vy nng
lng ti c mt bin ch cn khong 50% ca nng lng nhn c nh ca kh
quyn. Mt phn ca lng ny phn x tr li kh quyn t mt bin (hnh 3-12). Phn
phn x li ph thuc vo gc ti ca nh sng mt tri v tr nn rt ln khi gc mt
tri nh hn 5
0
so vi ng nm ngang. Trong mt ngy, nng lng bc x thc t
n mt bin ti bt k mt im l hm s ca gc ti nh sng mt tri, di ca
ban ngy v iu kin thi tit. Gc mt tri c xc nh theo thi gian ca nm, thi


gian ca ngy v v . Ti xch o, tng bc x hng ngy l hng s trong sut nm
nhng ti 50
0
v bc, s thay i bc x theo ma t 200 calo/cm
2
mi ngy trong
thng 1 n khong 950 calo/cm
2
mi ngy trong thng 6 (hnh v 3-11).

Hnh 3-11 S m t bc x mt tri qua kh quyn v mt bin v s cn i bc
x cho quang hp trong nc bin.

56



Hnh 3-12 (a) Ph bc x mt tri trc v sau khi i qua kh quyn, cho thy vng
PAR (bc x quang hp )hoc nh sng nhn thy. (b) Phn trm nh sng b phn x
t mt bin do gc ti ca mt tri, (c) Cc thay i tng bc x nhn c ti mt
bin theo kinh tuyn v ma Bc bn cu (ng ng mc l kinh tuyn).
2- Bc x ti mt bin v bc x quang hp thc t (PAR)
Trong nh sng mt tri ti mt bin c khong 50% thuc loi c bc sng
di hn 780 m. Bc x hng ngoi ny c hp th nhanh chng v chuyn sang
nhit su vi mt ca lp nc mt (Hnh 3-12). Bc x cc tm (< 380 m) ch
chim mt phn nh trong tng bc x v n cng thng b phn tn v hp th, ngoi
tr trong trng hp nc bin rt trong sch (Hnh 3-13). Cn li 50% bc x bao
gm nh sng nhn thy vi bc sng trong khong 400 -700 m xuyn ti su ln
hn. Cc bc x ny c tm quan trng c bit i vi ng vt c kh nng nhn thy
v cc bc x ny c cng bc sng m thc vt s dng cho quang hp. Nhng bc
sng ny gi l bc x quang hp thc t (PAR). Cng ln nht ca PAR khi mt
tri trc tip chiu vo l khong 0,6 cal/cm
2
mi pht. R rng gi tr ny s thay i
theo gc chiu ca mt tri v n gim xung ti 0 khi mt tri nm ng chn tri.
57


3- S tn tht nh sng trong bin l mt hm s ca chiu di bc sng
Khi nh sng i qua nc, n va b phn tn va b hp th. Cc bc sng
khc nhau ca nh sng nhn thy c th xuyn ti cc su khc nhau. nh sng
nhanh chng b hp th, ch khong 1% c gi li su 10 m trong nc bin rt
sch. nh sng xanh xuyn xung su nht, vi khong 1% cn li su 150 m
trong trng hp nc bin trong.
Quan h gia cng nh sng vi su nc bin biu th theo dng hm s
m. H s tiu hao nh sng k biu th s yu i ca nh sng. H s tiu hao nh sng
ca nc bin c th c tnh t cc quan trc, o c bng my bc x thp trong
nc bin v c tnh theo phng trnh (3-1), trong I
0
l bc x b mt v I
D
l
bc x ti su tnh ton, d l su (m):

d
I I
k
D e e
) ( log ) ( log
0

=
(3-1)
H s tiu hao nh sng k ph thuc vo chiu di bc sng, n khong 0,035/
m i vi nh sng xanh, nhng vi nh sng th n l 0,140/m. Mc d vy, nu
nhiu phn t c mt trong nc th nh sng xanh b phn tn nhiu hn nh sng .
iu ny s nh hng n quang ph mu ca nh sng di bin, lm dch chuyn
bc sng xuyn su nht sang mu xanh. H s tiu hao nh sng cng b nh hng
bi lng vt cht mang mu ho tan trong nc bin v lng cht dip lc cha trong
thc vt tri ni v thc vt bin. Trong nc bin sch nht vng nhit i, cc loi
c bin su c th nhn thy nh sng xuyn su ti hn 1000 m. Trong nhng vng
nc c ven b s pht tn v hp th nh sng tng ln do s c mt ca nhiu ph
sa v s lng ln cc thc vt tri ni, nh sng khng th ti su 20 m.

Hnh 3-13: Kh nng xuyn ca nh sng vi cc bc sng khc nhau vo nc bin.
Cc ng biu th su xuyn ti i vi 10% v 1% mc nh sng b mt nc.
58


C ba vng sinh thi khc nhau theo chiu thng ng ca nc bin c xc
nh bng s xm nhp tng i ca nh sng vo nc bin. Vng nng nht c
gi l vng sinh trng, n c nh ngha l vng c nh sng gip cho thc vt
pht trin.
Lng nh sng cn cho sn lng quang hp cn bng tn tht do h hp
trong thc vt c gi l cng nh sng b p. su m ti sn lng
quang hp c cn bng bi h hp ca thc vt c gi l su b p v n xc
nh bin di ca vng sinh trng. Nh vy vng sinh trng ko di t mt nc
n su ch vi mt khu vc nc c ven b v ti su ln nht khong 150 m
nhng vng bin nhit i nc trong. Ti bt k khu vc no su b p D
C
c
th c tnh bng cng thc:

k
I I
D
c e e
c
) ( log ) ( log
0

=
(3-2)
Trong , bc x b mt I
0
c o c trc tip v h s k c tnh t phng
trnh (3-1) vi gi thit bc sng 550 m. Gi tr cng nh sng b p Ic c
tnh bng thc nghim v ni chung thay i trong khong 0,001 v 0,01 cal/cm mi
pht, n ph thuc vo loi to chim u th trong vng.
Di vng sinh trng l vng nh sng m nht, y c c v mt vi ng
vt khng xng sng c kh nng nhn thy, nhng nh sng khng cho quang hp
dng (tc l lng tn tht vt cht thc vt qua h hp vt qu sn lng quang hp
ca cy trong 24 gi). D vy, thc vt ph du chm xung t vng sinh trng cng
c mt vng ny.
Vng su nht v ln nht trong i dng l vng ti thiu sng, n ko di t
pha di vng nh sng m nht cho n y i dng. Ti vng ny nh sng qu
yu n ni khng th nhn bit h sinh thi no. Vng rng ln ny khng gip g cho
cuc sng thc vt v n c chuyn tip theo khng gian t mt xch u tin trong
chui thc n ca bin.
3.2.2 Nhit (Temperature)
1- Nhit mt bin ton cu
Nhit nc bin c iu tit ch yu l do nng lng mt tri a ti v
s xo trn ca cc ngun nc khc. Ti nhng v thp, c s lu gi nng lng
mt tri ca tri t, nhng cc v cao th tri t li mt nhit. V th kt qu
l c s bin i nhit theo v ca b mt nc bin. S chuyn ti nng lng
theo b mt v chiu su l rt ln, v kh nng tip th nhit ln ca nc cho php
chuyn ti nng lng dc theo cc v . Tuy vy, nhit nc vng cc thp hn
0
O
C quanh nm, trong khi nhit nc vng nhit i hu nh u trn 25
0
C. S
hn ch cc b ca s lu thng nc trong cc h v m ph to ra s gia tng nhit
, i khi ln ti 35
0
C v cao hn.
59


S thay i nhit theo ma ln nht vng v trung bnh. Mc d nhit
ti cc v cao l nh, nhng thay i theo ma li nh nht. Nhit hu nh khng
i theo ma v m quanh nm l c im in hnh ca cc vng bin nhit i.
Thay i nhit theo ma vng Bc i Ty Dng nhiu hn cc i dng
khc l nh c nh hng ln ca cc h thng thi tit lc a. Ti vng bin ven b
Bc Caroina nhit thay i trong khong t 3 n 30
O
C. Ngc li, vi kh hu n
ho ca Thi Bnh Dng s thay i hng nm ch nh hn 5
0
C dc theo b bin
California, v nhit cao nht ma h thp hn nhiu so vi cc v i Ty
Dng. Nhng vng nc b hn ch lu thng v su nh nh a Trung Hi c s
thay i nhit hng nm ln hn i dng m lin k.

Hnh 3-14a Thay i nhit nc mt bin theo v i Ty Dng, Thi Bnh
Dng v n dng (Ngun: Anikouchine and Sternberg, 1973.)

i khi tin ch r cc vng a sinh thi ngi ta da vo nhit mt nc
bin. Cc vng bin c gii hn ca nhit mt nc bin trung bnh nm nh sau:
Nhit i 25
0
C
Cn nhit i 15
0
C
n i 5
0
C (gii hn pha bc)
2
0
C (gii hn pha nam)
Vng cc < 0 - 5
0
C hoc 5
0
C
Cc vng n i c hai bn cu c c trng bi s xo trn nc ca vng
cn nhit i v cn cc, chng c min thay i nhit hng nm ln nht. Mc d
c nhiu c gng gn gii hn v cho cc vng nhit ny nhng c rt t hoc
khng ng k v mt sinh thi trong mi trng bin ni cc dng hi lu y nc
c nhit khc nhau ra khi ngun ca chng v nhit nc thay i rt
60


chm thng qua xo trn. Trong cc qun x sng bin, cc ranh gii ng vt tun
theo cc ng ng nhit no (cc ng c cng nhit ), hoc c m t chnh
xc hn bng s kt hp nhit v mn dng xc nh cc th nc c bit.

Hnh 3-14b Nhit thay i trong bin (pha phi) v trong t lin (pha tri)
2- Phn b theo chiu ng - s phn tng
Trong nc bin, xo trn ri do gi v sng vn chuyn nhit t b mt xung
pha di . Ti cc v thp v trung bnh, iu ny to nn lp xo trn b mt vi
nhit hu nh ng nht t vi mt n hng trm mt chiu su. Bn di lp xo
trn ny, ti su 200 300 m i dng nhit bt u gim nhanh chng ti
khong 1000 m. Lp nc c s bin i nhit nhanh nht c gi l lp d nhit
theo ma, seasonal thermoclines. S khc nhau ca nhit qua lp ny c th ln
ti 20
0
C . Lp d nhit thng xuyn ny trng vi s thay i khi lng ring mt
nc gia nc b mt c khi lng ring thp, m v nc di y c khi lng
ring cao, lnh. Vng c s thay i khi lng ring ln c gi l vng
61


62
pycnocline, n hot ng nh mt vt cn i vi s lu thng nc theo chiu thng
ng, v th n cng nh hng n phn b thng ng ca cc cht ho hc ng vai
tr sinh hc ca bin. Cc bin i ln ca nhit v khi lng ring nc c th
cn tr s di chuyn thng ng ca cc ng vt.
Nhit gim dn pha di tng d nhit thng xuyn. S phn tng nhit
ca i dng c biu th hnh 3-15. Trong hu ht cc vng ca i dng, nhit
nc su 2000 3000 m khng bao gi vt qu 4
0
C bt chp c v . Ti cc
su ln hn, nhit h xung trong khong 0
0
C v 3
0
C. Nhit ca nc su ti
xch o cao hn khong vi so vi nc su vng cc, ngoi tr mt s vng cc
b no ni nhit nc y bin c th cao hn do hot ng a nhit.
Trong vng kh hu n i, thay i d thng nhit theo ma (seasonal
thermoclines) c to ra trong lp mt trong sut ma h. Kt qu ny l do bc x
mt tri tng ln dn n tng nhit b mt khi t gi. V th c t s xo trn ri
tng di chuyn nhit xung di v s phn tng nhit c hnh thnh trong lp
nc st mt. Hin tng ny ko di cho ti ma thu khi nc mt b lnh i v gi
tng ln to nn hin tng ri ng k xo trn cc lp nc trn v ph v
thermocline.
Kh nng sinh l phn ng vi nhit mi trng ng vai tr ln trong xc
nh gii hn phn b ca cc sinh vt bin. Phn ln cc ng vt bin (nh ng vt
khng xng sng v c) l cc loi bin i nhit vi s thay i nhit c th
gn vi nhit ca nc xung quanh, nhng cc ng vt bin c v li l loi
khng bin i nhit v gi c nhit c th khng i. Cc ng vt c th tn
ti trong cc mi trng vi min bin i nhit rng c gi l eurythermic.
Nhng loi nh vy thng c min phn b rng hoc chng sng trong cc vng c
nhit giao ng nh cc vng triu n i. Cc loi ny b hn ch i vi cc gii
hn nhit hp v c gi l loi t thay i nhit . Chng gm cc nhm nh
san h (yu cu ti thiu nhit l 20
0
C) cng nh cc loi b hn ch i vi nc
lnh. Phm vi a l ca cc loi t thay i nhit -lnh l rt rng, v d mt s
loi c tm thy vng nc nng ca Bc cc cng c mt su 2000 3000 m
cc vng xch o ni c nhit lnh tng t chim u th.



Hnh 3-15: Lc tng qut ho nhit vng nhit n ho. ng nt lin biu
th trng thi nhit ma ng vi lp mt trn ln c nhit ng nht nm trn
nhng im nng c nh. ng nt ri biu th s hnh thnh nhng im nng theo
ma pht trin lp nc mt vo ma xun v ma h, khi m bc x mt tri tng
ln, tri m hn v t gi.

Hnh 3-16: Biu m t cc tng nhit ch yu ca cc i dng v nhit trung
bnh ca chng ti xch o

63


3.2.3 mn (Salinity)
1- Cu thnh ca nc bin
mn l thc o cc cht rn v c ho tan trong nc bin, n l s gam
cc cht rn ho tan trong mt 1000 gam nc bin, biu th bng phn ngn (ppt hoc
%o). Nhiu nguyn t trong nc bin (Na, Cl, Sr) c tm thy c t l gn nh
khng i ngay c khi tng mui thay i t ni ny n ni khc. Tnh bt bin
ny tr nn r rng vi s gia tng thi gian lu li ca hu ht cc nguyn t trong
nc bin. Thi gian lu l thi gian trung bnh m mt n v khi lng vt cht tiu
tn trong bin trc khi n lng ng hoc chuyn ti cc lc a. Trong bin, cc
nguyn t c tm thy vi cc t l khng i c mt thi gian lu c mt triu
nm, ngc vi thi gian xo trn ca nc bin khong vi ngn nm. Cc nguyn t
khc c thi gian lu rt ngn. V d nh Nit v lu hunh, chng c to ra v tiu
th trong qu trnh sinh hc v cn i t ch ny n ch khc.
Nc bin l mt dung dch rt phc tp. Cc cht tan trong bao gm c cht
v c (bao gm c kh ho tan) v cc phn t hu c hoc cc sinh vt sng tri ni.
Cht ho tan v c vo i dng ch yu l do dng chy sng ngi, nhng cng c
th qua ma. Cc nguyn t chnh trong nc bin c nng ln hn 100 phn triu
(ppm). Chng gm clo, natri, magi, lu hunh, can xi, kali. Cc nguyn t chim u
th l clo v natri. V th khi nc bin b bc hi, mui clo natri cn li rt nhiu. Cc
nguyn t ph l nhng nguyn t c mt vi nng trong khong t 1 n 100 ppm.
Chng gm brom, cacbon, stronti, bo, silic v flo. Cc nguyn t vi lng c nng
khong 1 ppm bao gm nit, pht pho v st. Mt s ln cc nguyn t xut hin vi
khi lng nh v nng c tnh theo phn t (ppb).

Hnh 3-17: S thay i theo v ca mn b mt ca i dng m (ng xanh).
S cn bng gia s bc hi v lng ma c biu din trn ng mu en. (Ngun
Sverdrup et al., 1942).

64


T l clo so vi cc yu t khc trong nc bin c bn l khng i cho d c
nhng thay i hon ton v mn. Thc t ny cho php tnh tng mn bng vic
o lng clo trong nc bin. clo l tng nng ca clo trn 1000 ml nc bin
(tc biu th bng phn ngn). mn xp x 1,81 ln clo. Vic chun ho hc
mu nc v cc ion clo t nc bin c th c dng tnh tng mn. V dung
dch mui to nn dn in, nn cng c th dng dn in tnh mn thng
qua cc hiu chnh nhit . Mc d n ch chnh xc ti khong 2%o, my o khc x
quang hc cng c th dng o mn v s tng nng mui s lm tng khc
x nh sng.
2- mn
Trong i dng, mn thay i t 33 n 37%o. Ti nhng im nht nh,
mn thay i do cn bng pha long (nc sng chy vo, ma, cc mch nc
ngm) v qu trnh tp trung (bc hi, hnh thnh bng bin). S thay i theo v
trong cn bng ca ma v bc hi to nn mn ln nht 30
0
v bc v nam v
nh nht ti xch o v mn gim xung nhng v cao (Hnh 3-15). Hu ht
cc bin ven b khc vi cc vng lin i dng do s lu thng hn ch ca chng
v cc yu t a phng nh ma nhiu, dng chy t sng hoc bc hi. Bin Bantic
c mt lng dng chy ln v s lu thng vi i Ty Dng b hn ch nn kt qu
l mn trong bin Bantc nh hn khong 5%o. Ngc li, a Trung Hi c lng
bc hi qu ln so vi ma v dng chy nn mn ca n thng ln hn 36%o, n
nh hng ti bin Bc i Ty Dng bn cnh.

Hnh 3-18: S phn b mn b mt trung bnh (S) theo s khc nhau gia lng
ma v lng bc hi trung bnh (E-P) ti nhng v khc nhau
65



Cc gi tr mn gn nh tun theo ng cong i vi bc hi tr i ma nh
hnh 3-18. Ch rng cc gi tr mn cao nht l v 20
)
30
0
c hai bn cu,
trong cc vng c lng bc hi ln v ma nh. mn thp l cc vng cc, ni
c ma ln v bng tan, v cc vng chu nh hng ca nc vng cc.
3.2.4 Khi lng ring nc bin (Density)
1- Khi lng ring nc bin l mt hm s ca mn v nhit
Khi lng ring nc bin (khi lng trn mt n v th tch) ph thuc ch
yu vo nhit v mn (t ph thuc p sut thy tnh). Khi mn tng th khi
lng ring tng, khi nhit tng th khi lng ring gim. mn v nhit l
cc bin mang tnh vt l c lp, nhng nh chng ta bit chng khng c phn
b ngu nhin trong i dng.
Cc c trng mn v nhit ca cc khi nc bin c xc nh ti b
mt bin, nhng khi mt khi nc khng tip xc vi kh quyn, cc c trng vt l
ca n s thay i rt chm v rt nh thng qua s xo trn vi cc khi nc bn
cnh c cc c trng khc nhau. iu ny c ngha l mc d qua cc khi nc
chuyn ng theo c chiu ng v chiu ngang trong bin, mi khi nc c th c
tn ti theo khong cch nh s kt hp mn v nhit ca n. Cc khi nc mi
cui cng cng c hnh thnh nh s xo trn nc t cc ngun khc nhau v
chng hnh thnh cc du hiu nhit v mn ca bn thn chng biu th lng nc
xo trn.

Hnh 3-19: Biu T-S biu din mi quan h gia nhit (T), mn (S) v khi
lng ring. n gin, cc ng ng khi lng ring c biu din di dng
cc ng c cng gi tr
t
v
t
= (khi lng ring - 1) x 1000. V vy khi lng
ring l 1.02781 g cm
-1
c
t
= 27.81
66


67
Cc lp nc pha trn di chuyn ngang nh cc dng chy mt hnh thnh bi
cc h thng gi. Nhng chuyn ng thng ng ph thuc mt phn vo s thay i
nhit v mn. Hnh v 3-19 biu th mi quan h gia nhit , mn v khi
lng ring. Lu rng bn thn khi lng ring khng th dng nh ngha khi
nc v cc t hp khc nhau ca nhit v mn c th to ra khi lng ring
nh nhau.
2- nh hng ca khi lng ring nc bin n s lu thng ton cu
Kh hu ton cu to nn s phn b mn v nhit trong cc tng mt ca
i dng. Nhng t hp c bit ca cc bin ny c hnh thnh trong cc ct nc
ln v d dng o c, cc c trng nhit - mn c th c dng xc nh
mt khi nc c th. Mi c trng ca cc th nc ny to nn mt loi mi trng
khc nhau v mi loi nh vy thch hp cho cc cng ng sinh vt c th.
Nhng khi nc su nht v khi lng ring cao nht xut pht ban u gn
Nam cc hoc gn Greenland v Iceland (Hnh 3-20). Trong sut ma ng cc v
cao, nc b mt tr nn lnh hn v v mt nc bin tip tc tng ln n im
bng, nn c s chm xung lin tc ca nc cho ti khi t ti im bng. Khi nc
bin ng bng mn nh hn nc bin bnh thng do mn ca nc c
nng ln v khi lng ring cng tng. Nc vng cc c khi lng ring rt cao
chm xung v chy v pha xch o (Hnh 3-20) su trung bnh (nc vng gia
Nam cc v vng nc su i Ty dng) hoc dc theo y bin (nc y Nam
cc). Nc y vng Nam cc trong trng hp c th thm nhp vo phn pha bc
ca i Ty dng v Thi Bnh Dng. Nc su cui cng s tr li b mt nh xo
trn do gi, v v vy c mt chu k lin tc nhng rt chm gia nc mt v nc
su (vi trm n vi ngn nm). Trong sut ma ng ti cc v n i nc lnh
i bin su tip tc chm xung v khng bao gi t n im ng bng. Nh vy
mt i dng cn li bng t do tr nhng vng bin nng gn b nh Vnh St.
Lawrence pha ng Canada. Ngc li, nc cc h nc ngt c th lnh n 4
0
C
(nhit c khi lng ring ln nht ca nc ngt) t mt n y. Vi qu trnh
lnh gia tng, lp nc mt trong cc h tr nn nh hn v ni to nn s lu thng
thng ng. Sau nc mt tip tc lnh n im bng v s hnh thnh bng gn
mt l kh d dng.



Hnh 3-20: Mt ct ngang i Ty Dng biu din s hnh thnh v di chuyn ca khi
nc trung gian v khi nc su v cc a cc. AABW: Khi nc y nam cc; AAIW:
Khi nc trung gian Nam cc; v NADW: Khi nc su Bc i Ty Dng

3.3 Cu hi tho lun

68

Chng 4
thc vt ph du
v nng sut s cp

Trong i dng c mt s lng khng l cc thc vt khc nhau gm thc vt
ni v cc loi to n bo. Chng c gi chung l thc vt ph du. Nh chng ta
bit trong bin thc vt ph du c vai tr rt quan trng. Thut ng ph du l mt thut
ng thng dng trong sinh thi hc. Cc sinh vt ph du c nng lng di chuyn hn
ch, nhng chng sinh sng ni c s chuyn ng mnh m ca nc.
4.1 s phn loi
4.1.1 Thc vt ph du (phytoplankton) l g?
Thc vt ph du c nh ngha l cc sinh vt tri ni t do trn bin c kh
nng quang hp. Thc t cc thc vt ph du gm nhiu loi sinh vt quang hp khc
nhau. Thc vt ph du phn ln thng tn ti hai nhm chim u th nht l to ct
(diatm) v to hnh roi (dinoflagellates), cc thc vt ph du nh hn gm
nanoplankton v picoplankton vi kch thc rt nh. Mt s thnh vin ca thc vt
ph du nh hnh 4-1.

Figure 4-1: Mt s loi thc vt ph du: (a) to c, (b) to roi, (c) coccolithophore,
and (d) the microflagellate Isochrysis.
4.1.2 Cc v d v thc vt ph du v cc c trng ca chng
1. To ct: To ct d phn bit so vi loi to roi v chng c bao bi mt hp
trong nh thy tinh v kh thy s di chuyn. Mi hp nh vy c hai van (hoc v to
ct), mt van khp vi ci kia. Phn sng ca to ct nm trong hp. To ct thuc
69

lp n bo vi kch thc t bo khong 2 1000 m v mt s loi to nn cc
chui ln hn hoc cc dng tp hp khc trong cc t bo n l c gn vi
nhau nh cc gai hoc cc si nhy. To ct c nhiu bin t k phn trng (khong
100 triu nm trc) v qua mt thi k a cht di, lng ng cc v to ct c to
nn bi lng y bin gi l bn to ct.
To ct chim u th trong cc thc vt ph du t vng n i n vng cc.
Chng xut hin c dng cc t bo n gin v dng chui. Mi t bo c bao bc
trong mt v silic vi hai van khp vi nhau nh mt hp nh. Lp v c th c ph
bng nhng lng gai hoc c trang tr vi chui nhng li lm phc tp. Nhng ch
lm ch l lin kt gia t bo v mi trng bn ngoi. Cc to ph du thng to trn
v i xng (hnh dng ging vng pho hoa nh) v c gi l to trung tm, khc
vi to khc c dng i xng song phng v thng sng trong bn ct.

Hnh 4-2 : Chu trnh sng ca to ct
To ct sinh sn bng cch chia i t bo, vi mt van cho mi t bo con v
s dng nh mt van ln hn. Mt van nh hn c hnh thnh sau . Sau mt s ln
chia t bo lin tip, kch thc t bo thng gim xung mt ngng thp hn, khi
cc giao t c hnh thnh. Cc to nh hn cng c th hnh thnh mt bo t
sinh trng, n m rng v ct b cc van, to thnh nhng van mi ln hn, v sau
tri qua nhng ln phn chia v tnh. Nhiu loi to c th hnh thnh cc bo t v
tnh, chng nh c y bin v c th sinh sn sau ny nh mt dng sinh vt ph
du. V sinh sn v tnh nn to c th tng quy m dn s rt nhanh chng vi tc
70

gp i vi th t 0,5 n 6 mi ngy. To rt phong ph v loi v c th. Nhng nm
gn y ngi ta pht hin ra rng to sinh ra mt c t gi l axit domoic, axit ny
c th gy t vong cho cc sinh vt khc. Phng php sinh sn thng thng ca to
ct l chia v tnh n gin m trong t bo to ra hai nhn, hai na tch bit v mi
na to ra mt t bo mi nm trong v (Hnh 4-2). Cch thc ny cho ra hai t bo
mi c kch thc gn nh nhau, mi t bo mi nhn c mt na bn trong ca v
to c v hi nh hn t bo c hnh thnh t van bn ngoi. S phn chia v tnh
c th dn ti s gia tng dn s nhanh chng di cc iu kin ti u. Du sao vi s
phn chia lp li c th c s gim bt v quy m ca mt s th h sau.
Hai loi to ct c bit n l dng lng chim v dng trn. To lng chim c
hnh dng thon di v hu ht l sinh vt y, nhng mt s t loi sinh vt ph du nh
Nitzschia c rt nhiu mt s vng. To trn c cc van to trn xung quanh mt
im v chng nhiu hn bnh thng trong sinh vt ph du, vi khong 1500 loi.
Mt s loi to c m t hnh 4-3.

Hnh 4.3: To ct. (a) mt dng chui c trng ca Chaetoceros Iaciniosus; (b)
Chui C. laciniosus dng bo t khng sinh trng; (c) T bo ang phn chia ca
chui Nitzschia Pungens; (d) Chui Thalassiosira sp phn bo; (e) hai hnh dng v
to ct; (f) Coscinodiscus wailesii, xem hnh sau; (g) Chui Chaetoceros socialis dng
tp hp thnh tp on; (h) Mt dng chui ca Asterionella Japonica, v (i)
Skeletonema costatum. (T l l mm)
71

Cc loi to ph du khng c bt c cu trc di ng no v thng khng
kh nng chuyn ng c lp. To ny biu th mt s thay i c ch lm chm qu
trnh chm lng bi n l c trng ca to ct v thc vt ph du khc li trong
nc mt c nh sng nhm thc hin quang hp. S hnh thnh tp on hoc chui
cng tng din tch b mt v chm lng chm. Hu ht cc loi to thc hin iu
chnh ion, trong nng ni ti ca cc ion gim xung tng ng vi nng ca
chng trong nc bin. Trong t nhin, s ri lon ca nc mt cng rt quan trng
trong vic duy tr thc vt ph du gn b mt nc ni c nhiu nh sng mt tri.
2- To hnh roi: To hnh roi l dng khng c nhn, thng chim u th trong thc
vt ph du vng nhit i v cn nhit i v cng c th chim u th cc thc vt
ph du ma thu v ma h ca cc khu vc n i v phng bc. To hnh roi l
nhm ln th hai, sau to ct. Chng c hai roi c dng chuyn ng trong nc.
Chng thiu xung bn ngoi nhng thng bao bc bng cc tm xenlulo ccbon
hydrat. Cc roi ngang c mc rnh v chia t bo thnh hai phn gn bng nhau.
Cc roi khc c nh hng vung gc vi cc roi ngang v ko di ti pha sau. C
th thng c ph bi mt chui cc tm xenlulo k nhau.
Cc loi to roi khc nhau bi dng cc ngun nng lng khc nhau. Ch c
mt s loi to roi hon ton l t dng, to ra cc cht hu c v nhn ton b nng
lng ca chng t quang hp. Cc loi khc thc hin sn xut d dng. Mt s khc
na l kt hp, va t dng va d dng. To roi c chia ra hai loi
(Desmophyceae v Dinophyceae ) nh hnh 4-4.
To roi c c trng bi s c mt ca nhim sc th tch li qua chu trnh t
bo. Ni chung to roi l nhng sinh vt nh, n l, him khi to thnh chui. Chng
sinh sn v tnh bng phn i t bo. Cng nh to ct, chng c kh nng sinh sn
ti vi ln mi ngy. Sinh sn gii tnh cng xut hin v to roi c th to ra cc giai
on khng sinh sn, hoc cc t bo nang chm xung y bin v c th c ti sinh
sau ny nh l cc dng sinh vt ph du. Mt s to roi thuc loi pht quang sinh hc,
v ni bp bng nhng vng nc nng. Mt s to roi khc c kh nng gy c t
vo nc bin. Nu cc to roi tr thnh tp on ln (2 8 triu t bo trong mt lt),
th nh hng tch lu ca tt c cc c t tit ra ny c th tc ng xu n cc sinh
vt khc v gy ra ci cht hng lot. Nhng tp hp ln nh vy ca to roi c gi
l thy triu v l nguyn nhn gy ra cht c v cc loi khng xng sng hng
lot trn phm vi cc b nhiu ni khc nhau trn bin. Thy triu c th sinh ra
do cc loi to Alexandrium, Pyrodinium v Gymmodinium, chng gy tc hi cho
thn kinh gi l saxitoxin, loi ny c hn gp 50 ln strychnine v 10.000 ln
cyanide. Thm ch khi vi mt nng rt nh, to roi c th gy c cho ng vt v
con ngi.

72


Hnh 4-4: Mt s to hnh roi c ln. (a) Hai hnh dng ca Protorentfum marinum;
(b) Hnh dng ca Prorocentfuln micans; (c) P. micans ang phn chia; (d)
ProlopwdMim aassipes; (e) Gymnodiftm dng rt ngn li; (f) Dinophysis acuta; v
(g) GonyaulaK ftVft (Tt c cc thanh t l trn tng ng vi 0.02 mm.)
Ngun gc ca thy triu ging nh nhiu hin tng n hoa t ngt ca
thc vt ph du, l mt hin tng cha c gii thch y . Thy triu thng
kt hp vi hin tng t bin v dinh dng chy vo bin t cc sng hoc do ra
tri t t lin. Nhng trn bo c th di chuyn nhng khu vc bo xc b vi trong
bn ct y bin to nn mt giai on thy triu .
3. Thc vt ph du khc:
Cc loi gia vi (ngnh hong ngc) l loi sinh vt n bo v chng rt quan
trng trong cc loi thc vt ph du ca vng bin nhit i. Chng c dng gn nh
hnh cu v c bao ph bng nhng chui cc tm canxi ccbonnat, hoc cc gai vi,
chng thng ph y bin su. Trong nhm ny cn c mt s lng ln cc dng
trn km kh nng bo v. Chng c th l thnh phn quan trng ca mt s tp on
thc vt ph du.
Sillicoflagellates (ngnh hong ngc) l loi n bo v hai roi; chng c
nhiu lp lc v mt b xng bn trong bng silic dioxit. Chng thng to nn tp
on t hn to ct nhng ni ting trong thc vt ph du ca Nam cc v sinh vt ph
du ca nhiu khu vc bin h khc.
Vi khun lam (Cyanobacteria), l mt thnh vin ca loi to lam (thng gi
l to lam mc d thc t chng khng phi l to), chng xut hin v chim u th
73

cc vng nc ven b t lu thng cng nh cc vng nc l. Loi c lng
(filamentous Trichodesmium) thng thy nhng vng bin m ngho dinh dng.
Cc dng n bo b nh (khong 1 m) thng thy khp ni trn bin v l thc n
cho ng vt ph du. Cc vi khun c hi c th c nh m, trong nit c
chuyn ho thnh NH
4
+
, hoc ion amoni v sau kt hp thnh cc a xit amino v
protein.
To lc (Green algae), (n thuc lp to lc, ngnh hong ngc) y l loi
him trong nc bin nhng chim u th trong thc vt ph du cc vng ca sng
v m ph ven bin, c bit l vo cui ma h v ma thu. Chng c th l hnh roi
v khng chuyn ng. Mt vi loi gy hn ch cho s bng pht ca thc vt ph du
kt hp vi nhim vng ven b. To lc rt quan trng trong cc sinh vt y v l
nhng tp on rong to chim u th nhng vng chu thy triu.
To roi vng ngc (Cryptomonad Flagellates) l cc thnh vin ca lp to
vng (thuc ngnh hong ngc), rt ph bin cc vng ca sng. Chng l loi duy
nht v c cc loi dip lc a v c, cng nh dng di truyn hp th nh sng cho
quang hp.

Hnh 4-5: To lam k sinh Tricbodesmium tbiebautii (do Edward Carpenter chp)
4.2 Cc phng php thu thp mu
Cc mu sinh vt ph du thng c thu thp bng cc loi li. C rt
nhiu loi li c thit k khc nhau, nhng tt c u c c trng l dng hnh nn
di vi mt li nh. Ming ca li thng rng vi ng knh khong 50 100 cm
v c m nh b phn vng cng c 2 dy iu khin (hnh v 4-6). ui ca li
nh v c buc cht bng l thp hoc nha nh lc sch v gi li cc th cn
thu thp. Sau khi ko ly mu, li c ra sch v a vo v tr thch hp thu tt
c nhng g cn li trong li.

74


Hnh 4-6 : Mt mu li thu thp sinh vt ph du
lc c hiu qu, li phi c ko rt chm, khng nhanh qu 1 - 1,5 knot.
Mt li dy biu th cn cao i vi dng nc qua li, nu ko qu nhanh th
li s to ra hin tng ri trong nc v s lm tri mt cc vt ra khi ming li.
Nhiu kiu li c gng gim bt iu ny bng cch gn thm phn cnh bum
thun nh l li Hensen (hnh 4-7), n c kh nng ngn lng nc vo li hiu
qu lc c cao hn. Trc y, si t c dng lm li thu mu, nay cc li
hin i hu ht c lm bng si nilon.
thu bt nhng sinh vt ph du ln th mt li tha s hiu qu hn nhng
li dy v n gi li t hn v cho php dng nc chy nhanh qua li. Omori v
Ikeda (1984) khuyn ngh dng mt li vi c khong 75% ln ca loi sinh vt
nh nht phi ly mu v ko vi tc 0,7 1,0 m/s.

Hnh 4-7 : Li Hensen

75

thu mu cc sinh vt ph du ln nh cc loi tm, i khi dng cc li ln
nh li nh bt c con (YFTs) (Southward, 1970), loi ny c mt li c 1 mm v
c ming vi ng 1-2 m. thu mu cc loi sinh vt tri ni t do, gn thm vo
khung ni mt li ht b mt nc (David, 1965).
m cc sinh vt ph du, c b phn m trc tip s sinh vt nh c mt
trong nc chy qua dng c thu mu. B m l mt ng gm cc in cc ni vi
xung quanh ghi nhn s thay i tnh trng sp xy ra khi cc i tng i qua cc
in cc. N c kh nng tnh ton c s lng v kch thc ca sinh vt.
Hu ht cc kho st v s lng c thc hin cc mu sinh vt ph du c
lc qua t mt khi lng nc cho trc, phng php ph thuc vo loi nghin cu.
i khi cn c tnh tng s lng ca tt c cc loi sinh vt ph du. Khi lng s
b c th c tnh rt n gin bng vic cho php mu lng ng trong mt ng o
c v c th tch trc tip t vch o. Cc quan trc th tch thay th l kh chnh xc
hn, cc tnh ton c th c thc hin bng vic cn trng lng t hoc kh.
Hu ht cc kho st chi tit l m trc tip. Cc sinh vt ln thng c s
lng t v c th c nht ring m. Cc sinh vt nh hn c th chia nh mu
gim s lng nhm m c chnh xc hn. Cc mu sinh vt nh c th ri ra
trn a thy tinh phng m bng li vung. i vi cc sinh vt rt nh c s
lng ln th s dng huyt tc k m cc t bo mu hoc dng thit b m phn
t.
4.3 sn phm s cp
4.3.1 Quang hp
Thc vt ph du c gi l nh sn xut s cp chim u th trong i dng,
n chuyn i cc cht v c (nit, pht pho) thnh cc hu c mi (lipid, protein)
bng qu trnh quang hp v nh n bt u chui thc n trong bin. Mt lot cc
m thc vt c hnh thnh nh quang hp qua thi gian v c coi l sn phm s
cp hay sn phm quang hp v n l c s ca hu ht cc sn phm ca bin.
Mc d phi qua cc bc khc nhau nhng cc phn ng ho hc i vi
quang hp c th tm tt chung nh sau:
quang hp
(yu cu nh sng mt tri)
6CO
2
+ 6H
2
O = C
6
H
12
O
6
+ 6O
2
carbon dioxide nc carbohydrate oxy
h hp
(yu cu nng lng chuyn ho)

76

Cacbon dioxide c to s dng dng ho tan t do CO
2
, hoc CO
2
hn ch
nh cc ion cacbonat hoc bicacbonat. Tng s cacbon dioxit (c 3 dng) trong nc
bin l khong 90 mg CO
2
trn mi lt nc, v nng ny l ln nn n khng
gii hn lng quang hp bi thc vt ph du. Loi sn phm ny bao gm s gim st
cacbon dioxit to ra cc sn phm hu c nng lng cao c gi l sn phm t
dng; cc sinh vt t dng khng i hi cc cht hu c nh l mt ngun nng
lng. Ch rng qu trnh ny khng ch to ra cacbohydrat thc vt m cn to ra
cc o xy t do (nhn c t cc phn t nc ch khng phi t cacbon dioxit). Qu
trnh ngc li l h hp, trong c mt phn ng xy ho ph v nng lng nng
lng lin kt ln ca cacbon hydrat v v vy n gii phng nng lng cn thit cho
s trao i cht. Tt c sinh vt (gm thc vt) tin hnh h hp. Trong khi quang hp
ch c th din ra trong ban ngy, h hp li xy ra c ngy v m.

Hnh 4-8 : (a) S hp thu quang ph ca cht dip lc-a, (b) S hp thu quang ph
ca nhng sc t fucoxanthin (xanthophyll), khi c thm phycocyanin v phycoerythrin
(phycobilins).
Nng lng mt tri c dng iu khin qu trnh quang hp v s chuyn
i nng lng bc x thnh nng lng ho hc ph thuc vo cc sc t quang hp
c bit thng c trong lp lc ca to. Sc t chim u th l cht dip lc a, nhng
cc cht dip lc b, c v d cng vi cc sc t ph (carotenes, xanthophylls, v
phycobilins) c nhiu loi, chng cng tham gia vo s chuyn i ny. Tt c cc
sc t tch cc quang hp ny hp th nh sng bc sng khong 400 700 m
77

(PAR), nhng mi sc t biu th mt ph hp th khc nhau. Hnh v 4-8a cho thy
quang ph hp th ca cht dip lc a, loi thng xut hin trong sc t, hp th ln
nht l bc sng quang ph mu (650-700 m) v xanh tm (450 m). Hnh v 4-8b
cho thy quang ph hp th ca mt s sc t ph. Chng thng chim u th
khong mu xanh l cy ca cht dip lc v v vy nhiu thc vt ph du xut hin
mu nu, vng hoc thm ch mu .
4.3.2 Sn phm s cp
Nng sut s cp (primary production):
Nng sut s cp c nh ngha l tc to cc cht hu c giu nng lng
t cc cht v c. Nng sut s cp thng c xem l ng ngha vi quang hp,
nhng n khng hon ton ng v mt lng nh ca sn phm s cp c th c to
ra nh cc vi khun tng hp ho hc. Du sao y nng sut s cp c xem xt
vi qu trnh quang hp.
Tng lng cht hu c c c nh trong qu trnh sn xut s cp c gi l
tng sn lng s cp (gross primary production). V mt s trong tng sn phm ny
b thc vt s dng thc hin qu trnh sng ca chng, thng gi l h hp, mt
lng nh hn cn cho chuyn ho hoc cho cc sinh vt khc bin. Sn lng thc
(net production) l khi lng ca tng sn phm cn li sau khi tn tht do h hp ca
thc vt v dng cho cc mc dinh dng khc.
i vi c tng sn lng s cp v sn lng thc, cc t l thng c biu
th bng s gam cacbon c nh trn mt n v din tch hay th tch nc bin trong
mt khong thi gian. V d, sn lng c th c tnh bng gam cac bon trn mt
mt vung trong mt nm (g C/m
2
/nm), iu ny biu th sn lng tch lu trn ton
b chiu su t mt nc n y ca vng sinh trng.
Mt s thut ng khc nh standing crop, c p dng cho cc sinh vt t
dng, l tng lng sinh khi ca sinh vt biu th trong mt th tch nc cho trc
ti thi gian cho. Nng lng s cp v standing crop c th thay i ng k trong
mt khong thi gian t vi ngy n mt nm. S thay i ny l kt qu ca nhiu
yu t nh hng trc tip v gin tip ti qu trnh quang hp t dng ca chng.
Nc c nh hng ln n kh nng thm nhp ca nh sng. Do nc hp th
nh sng nn nng lng nh sng b tn tht khi ti vng nc su v cui cng l
bin mt. i vi cc sinh vt quang hp, c ngha ti mt su no sinh vt ch
nhn c nng lng vi mt t l bng t l n dng cho qua trnh trao i cht ca
bn thn chng. Nu cc sinh vt i ti su hn, qu trnh h hp vn cn mt t l
nh trn, nhng vic gim st nh sng s khng cho qu trnh quang hp v v vy
c s tn tht nng lng thc s. su ti tc h hp ca cc sinh vt ch bng
tc quang hp c gi l su b p (compensation depth). trn su ny,
tc quang hp vt qu tc h hp v c sn lng cc bon thc hoc sn phm
s cp thc.
78

su b p thay i theo cc vng bin trn th gii bi v n ph thuc vo
trong ca nc bin. Khi nc bin trong hn, s thm nhp nh sng ln hn v
su b p cng su hn. Ni chung, su b p su hn nc trong ti vng bin
h v n nng hn vng ven b c nhiu vt cht trong nc.
V su b p thay i theo su thm nhp nh sng, nn c th xc nh
su b p bng cng nh sng. Cng b p l cng nh sng m ti
quang hp bng h hp. Mc d cng ny c hi khc i vi cc loi thc vt
ph du khc nhau, n gn bng su m ti cng b gim xung cn 1% ca
cng nh sng ti mt nc. su b p c th xp x su ng vi 1% nh
sng thm nhp ti.
Cn nh rng su b p thay i khng ch theo cc vng a l khc nhau
ca cc i dng trn th gii nh cp m n cn thay i t ngy ny sang
ngy khc v theo ma trong bt k vng a l no. N l kt qu ca nhng thay i
nh sng theo ma, theo trong v cc cht dinh dng ca nc.

Hnh 4-9: Nhng t bo i lu Langmuir gy ra bi gi.
4.4.3 o c nng sut s cp
Phng php truyn thng c dng o c nng sut s cp c gi l
phng php bnh ti sng. Trong phng php ny, dng hai bnh ging ht nhau.
Mt bnh hon ton trong sut, cn mt bnh c qut sn en hon ton. Ly cng
lng nc nh nhau t mu nc cn tnh ton nng sut. Nc cha cc sinh vt ph
du mt cch t nhin. Lng xy trong nc c thm vo trong cc bnh mu tch
bit thit lp nng xy ban u. Sau cc bnh c nt li v gn vi mt si
dy, v a tr li su thch hp trong bin (qu trnh ). Trong bnh ti khng c
quang hp nhng thc vt v ng vt ph du trong vn h hp v tiu th xy.
Trong bnh ny, lng xy ban u s gim v ng thc vt ph du h hp s dng
xy ho tan. Ngc li, trong bnh sng hin tng quang hp vn tip tc vt qu h
hp, v vy gi tr xy tng hn gi tr ban u v n khng thot khi bnh. Sau khi
79

hai bnh c trong trong khong thi gian tnh ton, hai bnh c mang ln, m nt
v ngi ta phn tch hm lng xy trong nc. Thng thng nng oxy c
phn tch theo phng php Winkler.
T l quang hp c suy ra t nng oxy sau khi v nng xy ban u
ca nc khi ly vo bnh. hiu vic c lm th no, cn thit phi xem xt
iu g xy ra trong mi bnh. Khi nc bin ban u c cho vo mi bnh, n
cha mt lng xy c sn no (gi l nng xy ban u O
1
). Trong bnh sng
xy c dng cho cc sinh vt h hp. ng thi thc vt to ra nhiu xy thng qua
quang hp. Nng xy cui cng trong bnh sng l kt qu ca xy c b sung
do quang hp cng vi nng xy ban u tr i xy s dng cho h hp. V chng
ta bit lng xy ban u, tnh tng lng xy to ra do quang hp, chng ta cn
bit h hp tiu hao bao nhiu xy. Nu khng c h hp m ch quang hp, trong
bnh sng s tng ln ca xy s ln hn gi tr quan trc c. Trong bnh en, ch
c h hp xy ra v nng xy b gim xung. V c hai bnh c cng dung tch v
cng c vi thi gian nh nhau, s gim xy trong bnh en l tng xy do h
hp. tm ra tng lng xy to ra do quang hp cn lm mt vi php cng tr n
gin. Th nht, nng xy ca nc (trc khi ) c tr i t nng xy cui
cng trong bnh sng, v trong bnh en nng xy cui cng c tr i t nng
ban u. Gi tr ca phn tr i trong trng hp bnh sng l quang hp cng ng
thc, hoc lng xy quang hp vt qua h hp ca c sinh vt t dng v d
dng. N c gi l sn phm mi (new production). Gi tr t bnh en ch n
thun l h hp ca sinh vt. Quang hp tng cng (gross photosysthesis) nhn c
bng vic thm lng xy do h hp vo quang hp cng ng thc, hoc tr i lng
bnh en t bnh sng. N l s o ca tng xy quang hp xy ra trong sut thi
gian . C th chuyn cc t l sn phm xy thnh t l cac bon dioxid ng ho
bng vic s dng h s chuyn i. H s ny ph thuc vo cht hu c c to ra
(cacbon hydrat, lipid, hoc protein), nhng thng ly bnh qun l 1,2 1,4.
Mt phng php khc tnh nng sut s cp l phng php
14
C. Phng
php ny thng c dng tnh nng sut s cp trong khong 30-35 nm qua.
Trong , phng x
14
C c thm vo bnh cha nc bin c thc vt ph du.
14
C
thng c coi nh l H
14
CO
3-
(bicacbonat). Mt lng bit ca H
14
CO
3-
c b
sung v bnh c trong mt thi gian ti su thch hp di bin. Ti cui giai
on , cc bnh c ly ln v nc c lc qua cc thit b lc tt thu li tt c
cc sinh vt ph du. Lng phng x trong cc thit b lc c o c bng mt thit
b m. Lng
14
C xut hin trong mng lc t l vi nng sut. hiu chnh hp th
khng quang hp ca
14
C, mt bnh en thng c dng vi bnh sng. Cch tnh cc
bon hp th t phng trnh sau y:

(
14
C trong cc phn t thit b lc x
C hp th = cc bon v c c sn x 1,05) (4-1)
Tng
14
C c b sung
80


Cac bon v c c sn trong mu thng c o c bng vic chun kim.
Trong hu ht nc bin c mn hn 30 psu, tng cc bon v c c sn khong 25
mg trong 1 lit nc. H s 1,05 trong phng trnh hiu chnh cho s khc bit thc vt
ph du gia
14
C v
12
C, hp th
14
C chm hn.
C hai phng php u c sai s v u da trn cc gi thit khc nhau. Trong
phng php bnh en- trng, gi thit l cc iu kin trong c hai bnh l tng t,
ch tr nh sng. iu ny c th gy ra sai s. S pht trin ca vi khun lm tng
thm do s thm nhp ca b mt; V vy s lng vi khun tng ln c th nh hng
ti mc xy trong cc bnh. Vn ny thng xy ra i vi c hai phng php
14
C
v bnh sng ti. Mt vn khc i vi c hai phng php l cc sinh vt ph du
trong bnh khng hot ng bnh thng, v iu ny c th c phn nh trong t s
quang hp ca chng. i vi phng php bnh sng ti, mt vn nghim trng l
tnh khng nhy cm tng i khi nng thc vt ph du thp hoc khi nc b
nhim hay c cha nng vi khun cao. Vn khc i vi phng php
14
C l c
th c s ph v trong thit b lc
14
C r r ra ngoi,
Gn y, nhiu phng php khc nhau c dng nh gi cacbon dioxit
tng ln hoc xy b tiu hao cho thy rng phng php
14
C nh gi khng ng
nng sut s cp, c bit l trong trng hp nc c hm lng dinh dng thp v
mt thc vt ph du thp. Mt phn l do vn k thut (Peterson, 1980). Phn
khc c th l do nhim t cc kim loi kt hp vi cc mu bnh v cc dung dch
cung cp
14
C. V nhng thiu st ny m mt phng php mi v nh gi nng sut
s cp v ang c th nghim bao gm vic s dng nh gi quang hc quy m
ln thng qua nh v tinh v phng php quang- sinh hc ca vic o c quang hp,
c bit cc dng c o flo. Trong mt s trng hp, cc thit b ny o sc kho
sinh l hc ca thc vt ph du. Dng c c s dng trong cc nghin cu ny l
my hunh quang iu chnh ln nhp xung (PAM). Cn mt dng c khc, my
hunh quang t s nhc li nhanh (FRR) o t s quang hp tc thi.
Nng sut s cp thc (NPP) ca c hai qun x trn cn v di bin hin nay
c tnh t bc x mt tri hiu qu hp th quang hc (APAR) v e - hiu sut s
dng nh sng bnh qun theo phng trnh sau:
NPP = APAR * e (4-2)
i vi c mi trng nc bin v trn cn, APAR c xc nh t cc quan
trc v tinh thng qua vic o cht dip lc (trong mi trng bin) v xanh ca
thc vt (trn cn). Gi tr e khng th o c chnh xc t v tinh v phi c xc
nh bng cc quan trc thc t hin trng.
4.3.4 ng cong P-I v nng sut s cp trong mi quan h vi dinh dng
v nh sng
V ton b h sinh thi ca cc i dng ph thuc vo nng lng cho hot
ng quang hp ca thc vt ph du tng mt kh mng ca bin, ni c chiu
81

sng, nn cn phi hiu r cc iu kin ti nng sut ca chng c tng cng
hoc hn ch. Cc yu t quan trng nht l nh sng, nhit , nng dinh dng,
t v nc. Nng sut cy trn cn ch nh hng bi hai yu t l nh sng v cung
cp dinh dng, nhng trong bin th chng li b hn ch ng k. Do thc vt ph du
l lng trong nc bin chu chi phi bi cc lc lm chuyn ng ca nc v v nh
sng v dinh dng cng chu nh hng ca khi nc, nn cn xem xt mt nhn t
mi khng c trong t lin. Nhn t ny l nhn t hn hp c gi l thy vn hc
(hydrography), n bao gm tt c cc nhn t lm cho nc chuyn ng vng quanh
trong cc i dng, v d cc dng hi lu, cc khi nc tri ln v khuch tn. y
l s nh hng ln nhau ca ba nhn t ng sng, dinh dng v thy vn. Chng lm
hn ch nng sut thc vt ph du trong cc i dng v nhng vng a l khc nhau.
Lng nh sng (hay bc x mt tri) nh hng mnh m ti c khi lng v
tc quang hp. V vy quang hp xut hin trong mu t l vi cng nh sng
(xem hnh 4-10) ni quang hp tng ln vi s gia tng cng nh sng ti mt gi
tr ln nht (Pmax). Ti vng cng nh sng cao, c th c s gim ng k trong
quang hp (gi l s kim ch nh sng) n c gy ra do mt s phn ng sinh l
nh s hao ht ca lp lc trong nh sng trng.
im trn ng cong hnh 4-10 ti lng h hp cn bng mt cch chnh
xc vi lng quang hp c gi l im b p, im ny xut hin ti im cng
nh sng b p (Ic) nh l s nh du bin di ca vng nh sng. Thut ng
nng sut s cp tng cng (gross primary productivity (Pg) c dng m t
tng quang hp v khi nim nng sut s cp thc (net primary productivity (Pn) )
biu th quang hp tng cng tr i phn h hp.
ng cong trong hnh 4-10 c th c m t bng cc phng trnh ton hc
xp x vi hai chui phn ng c lp (biu th bng dc ban u P/I) l cc phn
ng rng buc ca quang hp v phn ng khc (Pmax) l cc phn ng ti. Cc
phng trnh n gin nht m t ng cong ti im Pmax (tc l vi s gim nh
sng) l:

] [
max
I K
] [I P
P
I
g
+
= (4-3a)
v

] [
] [
max
c
I I K
c
I I P
P
I
n
+

=
(4-3b)

82

y P
g
v P
n
l nng sut tng cng v nng sut thc; K
I
l hng s bn bo ho,
hay cng nh sng khi P = P
max
/2, cc gi tr K
1
thay i trong khong 5 n 25
E m
-2
s
-1
. [I] l lng xung nh sng PAR, v [I - Ic] l nh sng PAR xung quanh nh
hn cng nh sng b p Ic.
T cc phng trnh trn c th thy mt phn hi nh sng xc nh cho tt c
to pht trin trong cc iu kin vt l khng i v phn hi ny c th c tnh
bng hai hng s Pmax v K
I
. Thc t, cc loi khc nhau c cc gi tr Pmax v K
I

khc nhau v thm ch trong cng loi, phn hi quang hp ca mt t bo i vi nh
sng c th thay i theo thi gian. Ni chung dc ban u ca ng cong (P/I)
trong hnh 4-10 s thay i cng vi nhng thay i ca cc thng s mi trng nh
nng dinh dng v nhit , chng nh hng ti cc phn ng ti ca quang hp.
V cc loi khc nhau ca thc vt ph du phn ng khc nhau vi cc thay i ca bc
x b mt v ca cng nh sng. S thay i cc iu kin mi trng s thun li
cho cc loi khc nhau cc thi gian khc nhau v dn n s k tha ca cc loi
trong qun x.
i vi hu ht cc thc vt ph du, tc quang hp thay i theo cng
nh sng. Gn mt nc ni cng nh sng cao, qu trnh quang hp b suy gim
hu ht cc loi sinh vt (hnh 4-11a v 4-11b). iu ny l do c s hn ch quang
hp mc nh sng cao v s bo ho ca b phn quang hp, cho nn n khng th
tng tc . Cc loi khc nhau c cc ng cong tc quang hp khc nhau khi
c v vi cng cng nh sng v to ra cc cng nh sng ti u khc nhau
i vi quang hp cc i. iu ny c ngha ng k trong s din th theo ma.

Hnh 4-10 S phn ng ca quang hp (P) theo cng nh sng (I). /C S b cng
nh sng; K
I
l h s na bo ho hay cng nh sng khi quang hp bng 1/2
mc quang hp ti a (Pmax); Pg l tng lng quang hp v Pn l lng quang hp
thc.
83


Hnh 4-11a S quang hp ng vi cc ng cong cng nh sng cho 3 nhm
thc vt ph du. (T Biological Oceanographic Processes 3/e T. R. Parson et al. bn
quyn t @ 1984 Butterworth Heinemann. Ti bn bi Permission.)

Hnh 4-11b : S hn ch ca quang hp cc lp nc b mt th hin tng nng
sut ti cc cao trnh khc nhau ca ct nc. (Sau "Productivity, Definition &
Measurement" by E. Steeman-Nielsen t The Sea, Vol. 2, ed. M. N. Hill, 0 1963 John
Wiley & Sons.)
Cc cht dinh dng v c ch yu cn cho thc vt ph du pht trin v ti to
l Nit (nitrat, NO
3
-
; nitrit, NO
2
-
; or ammoniac, NH
4
+
) v Pht pho (nh phosphat,
PO
4
3-
). To ct v to roi cn cn mt lng silicat SiO
2
ng k. Thc vt ph du cng
cn mt lng nh cc cht dinh dng v c v hu c khc. Tt c cc cht dinh
dng ny c tm quan trng rt ln, mt phn l do chng c khi lng nh trong
nc bin. Chng l nhng yu t hn ch nng sut thc vt ph du tt c cc iu
kin. Cc i dng ca th gii c th c coi l nhng vng ngho dinh dng khi
so snh vi cc khu vc t lin. V d t canh tc nng nghip giu dinh dng cha
84

khong 0,5% nit tng t trn mt. Trong 1 m
3
t, lng nit ny cho sn xut
50 kg cht hu c kh. i vi thc vt trn t lin trong iu kin l tng c th to
ra vi kg cht hu c kh vt qu bn thn n cn cho mt nm trn 1 km
2
. Nh vy
t lin cc cy lu nm c th tip tc pht trin trong nhiu nm. V d, rng l biu
th ca s tch lu sn phm hu c trong nhiu nm. Tt nhin, cui cng tr khi nit
c ti to trong t, s cn kit Nit s lm hn ch sinh trng ca cy.
Mt s yu t c vai tr lm gim nng sut sinh khi. u tin, nh v d trc
gi thit rng hm lng nit ca nc bin l hng s trong ct nc. iu ny
khng hon ton ng. Cc lp nc trn mt thng c nng nh hn so vi cc
lp nc su hn. Tng t nh th, do nc hp th nh sng nn sn lng su
100 m s thp hn su 10 m. Quan trng hn, s tng s lng cc t bo thc vt
ph du c mt nh hng su sc. V qun th thc vt ph du pht trin lp pha
trn 100 m, cc loi cy c hp th nh sng nhiu hn. Kt qu l c t nh sng
xuyn ti c su hn. nh sng t hn c ngha l su b p bt u chuyn
ln trn v tr nn nng hn (hnh v 4-12).
Theo l thuyt, tng tc ln nhau ca cc yu t trn s lm gim nng sut s
cp. S gim nh su b p v hp th dinh dng l rt ln. Theo tnh ton, khi
qun x thc vt ph du t ti mt 2 g/m
3
th su b p 100 m ban u gim i
chng 3,5 m, v tt c nit c chuyn ho sang c th thc vt ph du. Sn lng cc
cht t dng cc i dng khng ng k so vi t lin. T l v sn lng gia
t lin v bin khc nhau khng ng k trn phm vi ton th gii, nhng v nng
dinh dng th khc rt ln v n bin i lin tc trong thi gian di.

Khng c thc vt S lng t
thc vt
S lng ln
thc vt
Hnh 4-12: S thay i ca su im b ng vi tng s lng thc vt ph du.
ng nt t ch su im b p.

85

Tuy nhin nu cc cht dinh dng b dng nhiu lp nc trn mt (c
chiu sng) th vn cn mt khi cha c khai thc trong khi nc pha di vng
sng. Bi v khi nc ny ln hn v dung tch so vi vng chiu sng, n nh mt h
cha dinh dng tr gip s gia tng ng k v sn lng. Nu h cha rng ln c
khai thc, th sn lng c th bn vng lu di v c bn.
ng tic l cc nhn t vt l thng ngn cn cc h cha ny trn hu ht
cc i dng th gii. Nc c cc khi lng ring khc nhau ph thuc vo nhit
v mn ca n. Nc lnh v mn th khi lng ring cao hn nc nng v t
mn. Pha di tng nh sng ca cc i dng nc b lnh v v th c khi lng
ring cao. Trong vng nhit i nc m v nn khi lng ring thp quanh nm.
Nhng vng n i, nc m trong ma h, c khi lng ring thp v lnh trong
ma ng. Ti cc vng cc, nc lnh v khi lng ring cao quang nm. Trong cc
vng nhit i s khc nhau v khi lng ring gia cc lp nc trn mt c nhit
cao v cc lp di lnh hn nn khng to ra s xo trn hai lp nc v v th cc
cht dinh dng khng th n c tng sng. Trong cc bin n i tnh trng tng
t chim u th trong ma h, nhng trong ma ng nhit ca hai khi nc tr
nn tng t v hin tng xo trn c th xy ra. Ti cc vng cc, khng c s khc
nhau ng k v nhit v s xo trn c th xy ra quang nm.
Mt iu khc cn bit hiu v b cha dinh dng, l c ch xo trn
hon ton. Trong khi nc c cng khi lng ring, cn mt s lc xo trn khi
nc. Lc c sn trn tt c cc i dng th gii l gi. Sc mnh ca gi thi trn
mt nc to nn lc xo trn. Khi hai khi nc tng t nhau v mt v nhit
th gi c vai tr to nn s xo trn ca cc khi nc v di chuyn nc giu dinh
dng ti cc vng c nh sng. Ti cc bin vng cc iu ny l c th xy ra bt
k thi gian no (khi bin khng b bng bao ph) v trong cc bin n i th ch xy
ra trong ma ng. Nhng s khc nhau ln v khi lng ring gia cc vng thiu
nh sng v sng trong sut nm ti cc vng bin nhit i v n i trong ma h
l ln, mc d khng c gi th chng vn xo trn cc lp nc. iu ny c
ngha l lp nc trn vn ngho dinh dng vng nhit i mi thi im trong
nm v vng n i l trong ma h tr khi cc iu kin thy vn vt qu s n
nh ca s khc bit v mt .
4.3.5 Nng sut thc vt ph du ton cu
Nng sut s cp ca thc vt ph du cc vng khc nhau ca i dng th
gii c c tnh t cc quan trc khc nhau v kt qu nh hnh v 4-13. Cc gi tr
thp i vi nng sut s cp (< 100 mg/cm
2
/ngy) thy cc vng xoy, ngc li cc
gi tr cao (>250 mg/cm
2
/ngy) thy cc vng bin n i v cc vng ven b. Cc
vng nc tri ven b c cc nng sut t ti 1000 mg/cm
2
/ngy. Tng cng, nng
sut s cp ca cc i dng th gii l khong 20 t tn cc bon mi nm. Con s
ny ngang vi sn phm pht pho ca thc vt trong t lin, nhng thnh phn sn
phm th rt khc nhau.
86

Trong cc h sinh thi t lin, cc nng sut rt cao thng c cc vng
din tch tng i nh vi sn lng rt cao. V d, nng sut s cp c tnh ca rng
ma nhit i l 3000 g/cm
2
/ngy, cao hn 10 ln nng sut thc vt ph du. Mt khc
nhiu khu vc t lin li l sa mc vi nng sut rt thp hoc khng ng k. Ngc
li, nng sut ca bin xut hin theo chiu ng bt k ni no trong vng sng
ca cc i dng (m bin chim hn 70% din tch mt t), thm ch vng cc.
N l s nh hng tch lu ca nng sut s cp bin trn ton cu lm tng thm vo
tng sn phm hng nm ca cc bon quang hp xp x vi t lin.
C s khc nhau theo ma v v v nng sut ca bin do s khc nhau v
nh sng v dinh dng c sn trong bin. Sn phm s cp c ng vt ph du s
dng cng khc nhau gia cc vng.

Hnh 4-13: Phn b nng sut s cp trong cc i dng (n v: mg C m
-2
/ngy)
Khi nng sut s cp tng ln, n lm tng ngun cung cp hin c ca thc
vt ph du. Trong sut thi k bng n vng ven b, ngun cung cp cht dip lc c
th tng t < 1 mg/m
3
n hn 20 mg/m
3
trong mt thi gian khong vi ngy. Ti mt
s vng, cc ng vt ph du c th n thc vt ph du ngang vi s pht trin ca thc
vt ph du, kt qu l d tng nng sut s cp rt kh nhn thy qua s gia tng ca
thc vt ph du. iu ny thy Bc Thi Bnh Dng, v khong 50
O
N (hnh 4-
14). Ngoi cc nh hng ven b, khng c thay i theo chiu thng ng v ngun
cung cp thc vt ph du trong nm, n l hng s khong 0,5 mg cht dip lc/m
3
.
Mc d nng sut s cp trong vng ny tng t ma ng < 50 mg/cm
2
/ngy ti hn
250 mg/cm
2
/ngy vo thng 7. Nng sut s cp vt qu c tiu th ca cc ng
vt ph du ti vng (hnh 4-14). Giai on thc vt ph du v ng vt ph du c s
87

quan h mt thit i vi cc sinh vt y bin su vng bc Thi Bnh Dng l khi
cc thc vt ph du nh chm xung vng nc su to ra thc n cho ng vt y.
Ngc li i Ty dng cng v , s bng n trong ma xun c c
trng bi s gia tng cht dip lc ln hng chc ln, t 0,1 n 1,0 mg/m
2
. Nng sut
s cp tng ln nh Bin Thi Bnh Dng, nhng ng vt ph du t hn. Do ch c
mt phn nh thc vt ph du b tiu th. Nhiu ni Bc i Ty Dng cng c s
bng n trong ma thu ca thc vt ph du, trong hnh 4-14 khi c nh th hai ca
sinh khi ng, thc vt ph du. Rt nhiu thc vt ph du khng b n Bc i Ty
Dng, phn hu bng n lng xung nc su v tr thnh ngun thc n cho ng
vt y bin.

Hnh 4-14: Tm tt chu trnh hng nm ca cc qun x thc vt ph du cc vng
khc nhau. (ng m l thay i sinh khi thc vt ph du, ng t l thay i
sinh khi ca ng vt ph du).
Hai chu trnh hng nm khc nhau ca ng, thc vt ph du c trnh by
hnh v 4-14. Mt chu trnh cho thy thnh phn bin bc cc, ni c nhp sng n
gin ca thc vt ph du xut hin ngay sau khi bng gi bin mt v tip theo l s
chm chp ca nhp sng n gin trong sinh khi ca thc vt ph du. Thi gian tr
trong phn ng ca ng vt ph du i vi thc n gia tng l do tc tng trng
tng i chm trong nc lnh. Trong cc mi trng nhit i th sinh khi ca ng
thc vt ph du cho thy khng c s thay i bn vng trong nm. D sao cc hot
ng ca bo c th lm thay i iu ny tr khi mi trng rt n nh n ni cc
thay i nh trong sinh khi ca sinh vt ph du c th xut hin khng u trong
nm. Ti cc vng nc m nhit i, bt k s gia tng no trong ngun cung cp
thc vt ph du u c nhanh chng tiu th bi s pht trin ca ng vt ph du.
88

Nng sut s cp thay i theo chiu su, v phn b theo chiu thng ng
ca thc vt ph du c th thay i theo ma. Ti nhng v n i, thc vt ph du
s c xo trn tt tng nc mt trong sut nhng thng ma ng v bt k s
quang hp no cng s tun th ng cong gim st nh sng, ngoi tr mt s hn
ch nh sng vng st mt nc. Do pht trin trong ma xun, nng sut s cp s
gia tng trong ngun cung cp thc vt ph du. Cui ma h khi h thng b ht ngun
dinh dng st mt nc, nng sut s cp ln nht s chuyn xung lp nc su hn.

Hnh 4-15 : Biu thay i theo su ng vi cc ma ca sinh khi qun th thc
vt ph du (S), Tc quang hp thc hng ngy (Pn) v nng cht dinh dng
(N) theo nhit tng nc. S (Vng k sm), n v thng l mg Chl-a m
-3
; Pn
(ng en), n v thng l mig C per mg Chl-a/ngy, N (ng sng), n v
thng l M nitrate.
Trong cc khi nc n nh (hu ht cc bin nhit i v cn nhit i),
phn b dinh dng theo chiu ng, nng sut s cp v cc s tng ng v cht
dip lc cho thy vo cui ma h nh hnh 4-15 v l c tnh ca khi nc qua
nhiu nm. Gi tr ln nht ca cht dip lc trong nc c th c thy bt k
ni no trong khong t 20 m n trn 100 m su v ph thuc vo s n nh ca
nc trong thi gian di. Di cc iu kin ny, vng ph dng thc t c chia ra
2 qun x theo chiu ng. Qun x pha trn b hn ch v dinh dng v c bn da
vo cc qu trnh sinh hc v ho hc ti to dinh dng trong vng. Qun x di y
b hn ch nh sng nhng n l vng giu dinh dng, thay i ln nht trong
nng dinh dng xut hin, v v vy cc ngun dinh dng b sung vo h thng l
t nc su hn. V mt s thc vt ph du v c di c dc theo hai qun x ny nn s
c mt chuyn i sinh hc gia hai mi trng tch bit theo chiu thng ng.

4.4 CU HI V THO LUN

89


Chng 5
ng vt ph du
5.1. Phng php phn loi
5.1.1 ng vt ph du l g?
Ni mt cch ngn gn, ng vt ph du (zooplankton) l loi ng vt sng l
lng trong nc v nh nc tri i. Ngc vi thc vt ph du ch c kh t sinh vt
quang hp, ng vt ph du cc k a dng, bao gm i din ca hu ht ng vt v
nhiu ngnh sinh vt nguyn sinh. C th chia ra lm hai nhm: (i)- ng vt ph du
thc s ln; v (ii)- ng vt ph du c nh (cc nh v siu nh). Chng bao gm cc
sinh vt n bo c kch thc rt b cho ti loi sa c ng knh ti vi mt. Mc
d tt c cc ng vt ph du u c kh nng di chuyn, nhng nh nh ngha
nu, khng c loi no trnh khi b dng nc cun i. Tt c cc loi ph du, c th
l ng vt v mt vi loi vi sinh u l loi d dng. iu ny c ngha l chng
cn cc cht nn hu c (ngc vi cht nn v c) nh l ngun nng lng ho hc
tng hp cc cht cn thit cho c th. Khng ging nh thc vt t dng bng
cch s dng ngun nng lng mt tri lm gim Carbon dioxit (CO
2
), ng vt hp
th carbon v cc cht ho hc thit yu khc thng qua qu trnh tiu ho cc cht
hu c. Cc loi ng vt khc nhau c cch tip nhn nng lng khc nhau: loi n
c th tiu th thc vt, loi n tht th ch n tht ng vt khc, cn loi n cht mn
th tiu th cc cht hu c thi ra. Tuy nhin, c nhiu loi ng vt n tp. ng vt
ph du thng c xp loi theo cch n ca chng.
5.1.2 Mt s th d v loi v cc c tnh ca chng
Trong s cc loi ng vt ph du c mt nhm ng vai tr quan trng hn cc
nhm khc. Lp chn cho (ph ngnh Crustacea, ngnh Arthropoda) bao gm cc
loi gip xc ni hon ton c kch thc rt nh xut hin kh ph bin trong s cc
loi ng vt ph du trong th gii i dng (Hnh 5-1). Cc ng vt b nh ny c
vai tr ht sc quan trng i vi cc h sinh thi bin v chng l mt trong cc loi
ng vt n c bc mt i dng. Chnh chng n cc loi thc vt ph du ln sng
trong nc, c bit cc vng ven bin v to nn cc lin kt gia cc sinh vt t
dng bc mt v cc loi n tht ln nh khc nhau.
Ngoi vic phn hng v v tr trong dy chuyn thc n, ng vt ph du cn
c th c chia nh ra na theo mi trng sng (loi sng i dng khc vi loi
sng ven b) v theo cch phn loi khc. Chng cng c chia ra hai loi ph
thuc vo thi gian sinh sng trong mi trng bin khi: (i)- ni vnh vin
(holoplankton) l sng ton b cuc i ca chng trong vng nc ; (ii)- ni theo
giai on (meroplankton) l nhng c dn tm thi ca cng ng ph du. Loi ni
90

tng giai on bao gm trng c v u trng (khi trng thnh loi ph du), bao
gm c giai on l u trng tri ni ca cc ng vt khng xng sng di nc
nh trai, c, a v sao bin.

Hnh 5-1 : Mt vi ng vt ph du: (a) mt loi tm, (b) B ru nhnh Podon, (c)
Mt loi thuc b Trng l, (d) mt loi trng tia, (e) sa lc, (f) su c lng, (g)
loi sa thuc lp Scyphozoa v (h) thc vt tri ni t do c l siphon

Hnh 5-2: ng vt chn cho. (a) ng vt chn cho in hnh v cc c im
ca cc c quan chnh (b) S chu trnh sng in hnh ca mt ng vt chn cho.
91

a)- Gip xc ph du (Crustacean Zooplankton): Cc loi gip xc to nn mt lp
ca ngnh Arthropoda, c cc c im ring bit sau: (1) phn xng to bi cht
kitin cng pha ngoi, c th un c v c kh nng ngn cch kh tt vi mi
trng bn ngoi, (2) phn ph c mt s on ni vi nhau v mc cn xng (chng
hn nh chn, ru). Cc loi gip xc c phn u bao gm ru, hm di, hm trn, v
thng c mt kp. Chng bao gm cua, tm, tm hm, tm nc ngt v mt g (loi
sng trn cn). Cc c im ng ch ca chng l tnh di ng, c phn v gip
xc bn ngoi v d nhn thy.
ng vt chn cho (Copepods), trong s cc loi ng vt ph du th loi
chn cho chim nhiu hn c, cc sinh vt ny c mt hu ht cc i dng v
vng ven bin. Gip xc thuc b chn cho cho ti nay l nhm gip xc ln nht
trong loi ng vt ph du. Chiu di ca chng t nh hn 1 mm cho ti vi mm. Nu
ni v khi lng, ng vt chn cho calanoid nhiu hn bt k mt nhm no khc.
Harpacticolds thng sng y trong giai on trng thnh, nhng u trng ca mt
s loi khc c th chim u th vng ca sng. Cc chi thuc h Lernacopodoida
k sinh trn c v ng vt c v sng bin. Chng ging nh su v c th nhy t
vt ch xung nc vi mt khong cch 30 cm.
Cc loi chn cho Calanoid thng c dng ng trn, thn gm c u, ngc
v bng. Chng bi nh nh nhp cc i ru pha trc v cc i chn phn
ngc pha sau. Chng c mt con mt nm chnh gia nhng khng phi mt kp
nh nhiu loi gip xc khc. loi Caralus, c khong 10 n 14 ngy, con ci ra
mt trng c khong 50 ci. u trng tri qua giai on sinh trng v lt xc trc
khi trng thnh.
Calanoid ch yu n thc vt ni, mt vi loi cht hu c v ng vt ph du
nh b hn. Chng pht hin thc n nh i ru nhy cm nm pha trc. Chng
gi cc ht thc n lng t mm nm ngay trn lng cng khoang hm trn. Trong
mi trng cht lng c s Reynold nh, khi tc dng chy nh v kch thc cc
vt th trong cht lng nh th nc chy ri qua mt ng vt chn cho v khng y
vo chn ca chng, m lc ny dng chy nht chim u th, chy bao quanh chn.
Chn cho s p bn cp chn y nc. Khi to ct tin gn, hm trn ca chn
cho vn ra v chp ly to ct, ch khng lc n qua cc lng t (nh lu nay ngi
ta vn tng). cc loi chn cho ln hn, s Reynolds khong bng 1 v c s ng
gp ca cc lc qun tnh. cc iu kin nh vy, hin tng lc bng lng t c th
xy ra. Chng phi pht hin s c mt ca to ct bng cc b phn nhy cm phn
ph (ru, chn) trc khi chp ly con mi.
Krill l cc chi ca h gip xc Euphausiidae ging nh tm, di thn ln
nht l 5 cm. Chng l loi ng vt ph du c mt nhiu nht bin Nam cc nhng
cng kh ph bin nhiu vng bin khc trn khp th gii, ni c ngun ti nguyn
phong ph, chng hn nh dng hi lu Benguela ca Chu Phi. Cn Nam cc,
chng l thc n ch yu ca cc loi c voi sng hm. Krill n thc vt ni v cc loi
ng vt ph du nh hn, nh ba cp chn phn u ngc lm nhim v nh ci gi.
92

Ging nh chn cho calanoid, krill c cc chn mc lng cng v lng mm c nhim
v bt thc n. Tinh trng di chuyn qua cung sinh tinh v trng c gi nh chic
gi phn ngc hoc ti trng nm bng.
Hu nh mi loi tm vy (euphausiids) u pht sng. Cht pht sng ni
bo v nm c quan pht sng, thng l phn u trn ca cung mt.
Ru nhnh (Cladocera) c vng nc ngt nhiu hn l vng nc mn,
nhng i khi cng kh ph bin vng ca sng. Thc n ca chng l cc ng vt
ph du khc.
Mt vi loi gip xc khc, nh tm cm, cng thc s l loi ph du nhng t
khi ni, chng ni vo ban m nhng ban ngy li sng di y. Cng c mt vi
loi nh Eutbemisto v Hyperia l ng vt ph du ni hon ton v sng nhiu ni
khp cc bin trn th gii.
b)- ng vt ni dng trong sut v mm:
Ngnh rut khoang hay sa c lng chm, lp Scyphozoa - thc s l sa -
c chia thnh mt s b khc nhau. Sa chuyn ng ln pha trc bng cch nhp
nhng co bp d (hnh 5-1g). Chng bt ng vt ph du bng cch chch xc tu vo
con mi vi cht keo dnh gi l t bo chm nga (nematocyst). Mt s t bo ny c
cha nc c v mt s loi ong bin (Cubomedusae) v loi khc nh hnh 5-1h gy
au v thm ch c th git cht ngi.
Sa c l siphon l mt qun th c chng v l nhm lng chm c nhiu
dng. Cc b phn khc nhau m nhn cc chc nng ring bit nh tm bt thc n,
bi v sinh . Loi c tn l Thu th Velella c kch thc nh nhng c mt phn
rt ging vi cnh bum c th n gi, chng sng rt ng c ngoi khi Thi
Bnh Dng v thng b dt vo b bin dc California ti Washinton. ng vt ny
c mt lp o bao quanh phn bong bng c cha kh gip cho con c thng bng khi
gi p vo cnh bum. Cnh bum l mt tm phng li dng cn gi tri theo
chiu gi. Khng ging nh nhng cnh bum ca cc con thuyn bum c hnh tam
gic cao hin i, bum ca Velella khng ng dng nguyn tc Bec-nu-li to sc
nng. Mt loi c lin h kh mt thit l Porpita dng nh cng c b phn to thng
bng di dng cc xc tu to trn, nhng khng c bum.
Cc loi sa lc l cc chi ca ngnh Ctenophora, thuc loi jelatin, gn nh
trong sut v c dng hnh trng. Sa lc khc cc loi ch chng c 8 hng l nm
dc pha ngoi, nhng mt s c xc tu di (hnh 5-1e). Chng l loi n tp v c
bit a n cc ng vt chn cho. Pleurobrachia bt chn cho bng cc xc tu xon
trn di v a xc tu ln pha ming. Mnemiopsis khng c xc tu di v cc lng
mao a con mi vo nhng ci khe ni con mi c gi li nh mt hng cc xc tu
ngn v sau cc xc tu ny a mi vo khoang mi, mt b phn dn thng ti
ming. C cu ly thc n ny cc k hiu qu v ngi ta tin rng chnh sa lc gy
ra s suy gim s u trng ca loi s, hn v tm cua. Cc giao t thng c phn
93

tn i trong nc v phi thai pht trin thnh u trng trng ging nh c trng
thnh. Sa lc c kh nng pht quang sinh hc mnh v pht sng nh nhng bng
n khi b quy ry.
Salps, Thallacea (ngnh Urochordata) l loi c bit i din cho cc ng vt
ph du bi t do. Khng ging nh h hng ca chng l loi bm tim bin sng
y, chng c cc ng nhp v xut cc u i din nhau trn c th. Salps c dng
ng trn ngn v sng n c. Pyrosoma, ngc li sng theo by n v c dng ng
trn di bt kn mt u. Cc qun th ny c th di ti 2 m v mi c th c ng xut
hng ra ngoi, ng nhp hng vo khoang trung tm. Salps gi thc vt ni v cc
ht nh trn mt mng lng nhy v n cc ht c ng knh t 1 m ti 1 mm. Mt
s loi salps ch yu l n u trng tm c.
Loi larvacea (ngnh Urochordata) l mt nhm ng vt ni geletin c bit
khc, n c chiu di thng ch c vi mm v c mt s c im ring in hnh cho
loi u trng bi c v. Loi ny to nn mt ngi nh v gn cht mnh vo hay
bn trong ngi nh . Nhp p ca n to nn mt dng chy qua "ngi nh".
Dng ny c lc qua mt li t mn tri pha trc by thc n. Cc ht c
chuyn vo ming v phn loi trong cung hng.
c)- Cc loi ng vt ph du khc :
Su c lng - Arrow worm (ngnh Chaetognatha), c dng nh qu ng li (di
4-10 cm), vi mt hoc hai cp vy bn cnh sn (hnh 5-1f). Chng bi nhanh nh
co bp nhanh cc bp tht nm dc trn thn. u c trang b nhng chic ngnh
bt mi, thc n a thch ca chng l cc loi ng vt ph du khc. Cc loi khc
nhau thng sng nhng vng nc c khi lng ring khc nhau, v c th phn
bit c loi sng ngoi khi vi loi sng vng ven b. Chng l loi lng tnh,
trng c th c phn tn trong nc hay bm vo cc vt ni ri n ra u trng v
ln ln.
Pteropods l cc loi c hon ton sng ni v bi bng mt b phn mc
ngang chn khc vi loi c sng st y. Nhiu ni Pteropods sinh si mnh, v v
chm xung y to nn mt lp trm tch gi l bn Pteropods.
Polychaetes ni, cng l ng vt ph du ni hon ton, c chn rt pht trin
(chi bn) v cc c quan cm gic (chng hn, loi Tomopteris).
4- Ph du hon ton v ph du tng giai on
a)- Ph du hon ton : C khong 3700 loi ph du ni hon ton i din cho nhiu
nhm ng vt khng xng sng khc nhau bit. Cc nhm thng xut hin
bin v gp vai tr quan trng trong qun x ph du s c m t di y.
ng vt ph du nh nht l sinh vt nguyn sinh n bo. Loi ny bao gm
rt nhiu loi trng roi, mt phn hoc hon ton l d dng. Cc trng roi d dng
ny n vi khun, to ct, cc trng roi khc, v cc ng vt nguyn sinh c lng hoc
do dng nc mang ti hoc do chnh chng bt c. Mt s loi c kh nng d
94

dng, mt s khc c cha lp lc lm nhim v t dng trong mt giai on no
. Loi d dng c bit n nhiu nht c dng mt qu cu trong sut vi ng
knh khong hn 1 mm. Noctiluca thng xut hin cc m ly rm rp gn b
bin, v n cc loi ng vt ph du nh hn (bao gm c trng tm, c) v to ct
cng cc loi thc vt ph du khc.
Mt nhm ng vt nguyn sinh hnh roi, thng c gi l zooffagellates,
gm tt c cc loi khng mu v hon ton l d dng. Tt c cc sinh vt trong
nhm ny thiu lp lc, sc t v nhiu loi n vi khun v cc cht thi ra. Mc d
rt nh (thng 2-5 m) nhng chng c kh nng sinh cao v do sinh si rt
mnh trong nhng iu kin sng thch hp. Tnh v s t bo, cc trng roi d dng
chim khong 20-80% trong s sinh vt ph du cc nh, v do vy chng c th l
ngun thc n ch yu cho ng vt ph du chuyn n cc sinh vt b nh.
Trong s cc amip bin c Foraminifera, l loi c v hay mai to bi canxi
v thng c nhiu khoang. Kch thc ca cc sinh vt ni khong 30 m ti vi
mm. Thc n gm cc vi khun, hay ng thc vt ni c nh c bt gi bi mt b
phn c bit mc ra t l ca v c gi l rhizopodia. Mc d c cha ti 40 loi
sinh vt ph du c bit (trong khi ng vt y l khong 4000 loi), cc loi c l
ni hon ton ny pht trin rt mnh, c bit vng khong t 40
o
v Bc n 40
o

v Nam v sng trong vng nc su khong 1000 m.
Loi Radiolaria c hnh cu, thuc ngnh ng vt nguyn sinh amip, vi nang
c l to bi silic. Kch thc ca cc sinh vt ny c ng knh khong t 50 m ti
vi mm, mt vi loi kt thnh nhng qun th jelatin t cc c th v di ti mt vi
mt. Radiolaria thng sng cc vng nc lnh v su.
Loi ph du c lng mao c mt tt c cc i dng v thng rt phong
ph. Tt c u s dng lng mao trong qu trnh vn ng, mt s c lng khc kiu
ming tm bt thc n. Nhng loi ny n sinh vt dng roi, cc loi to ct nh v
vi khun. Tintinnids to nn mt tiu nhm (hn 1000 loi) ng vt ph du c lng
mao. D c kch thc nh (khong 20 - 640 m), nhng tintinnids c vai tr sinh thi
rt ng k v chng c mt c nhng vng bin khi v ven b, thc n ch yu ca
chng l to ct ni cc nh v sinh vt hnh roi c kh nng quang hp. vng ben
b, tintinnids c th tiu th t 40-60% lng thc vt ph du. V chng cng chnh l
mi cho rt nhiu loi ng vt ni tng gia khc.
Sa (jellyfish), hay medusae l loi d thy nht, thng sng bin khi v
khu vc gn b. Mt s loi sa ni hon ton, nhng mt s khc c mt giai on
sng v tnh y trong chu k sinh trng, c loi sa l ph du ni theo giai on.
Mc d jellyfish thuc mt s nhm khc nhau trong ngnh sa c lng chm
(Cnidaria), nhng tt c u c mt t chc cu trc c bn v u l loi n tht, n
cc ng vt ph du bng cc xc tu c trang b cc t bo chm nga. ng knh
ca chng dao ng t ch vi milimt cho ti vi mt i vi loi Cyanea capillaia,
mt loi thng thy pha bc c s xc tu ln ti 800 hoc hn vi chiu di 30-60
95

m. Ngi ta bit kh r mt s loi sa c lng chm sng dng qun th, ging nh
loi c l siphon, trong nhiu b phn vi cc chc nng chuyn bit c hp nht
li to nn c quan tng th. Physalia hoc Portuguese man-of-war, l loi qun th
c bit n nhiu nht (Hnh 5-3a).

Hnh 5-3: Sa c l siphon (a) mt loi ni mt, xc tu di ti 10 cm; (b), mt loi sa
bi, di 10 cm. C hai loi u s dng cc xc tu ui bt mi.

Sa phin lc (Ctenophores) rt gn vi sa, nhng cu to ca chng cng
khc cho chng c xp vo mt ngnh ring. y l loi ng vt trong sut
ging nh Sa c lng chm, n l loi n tht, nhng khng c t bo chm nga nh
nhng h hng rt gn vi chng. Mt s Ctenophores nh loi mang bn
(Pleurobrachia) c cc cp xc tu di vi cht keo dnh bt mi; mt vi loi khc
(chng hn nh Bohnopsis) bt thc n bng cc thu c lng ming. Sa phin lc
ny c nh hng kh ln n lng c v chng n trng c v u trng c, chng
cng cnh tranh vi c con n cc ng vt ph du nh hn, chng hn nh loi
chn cho. Mt s loi sa phin lc nh Beroe khng c xc tu nhng li c nhng
ci ming rt to.
Lp hm t (Chaetognaths hay trng lng) l mt trong cc nhm ng vt
ph du n tht phong ph nht v c bit n nhiu nht. Nhng ng vt lng tnh
ny ch thy xut hin bin, di tng nc su n vi nghn mt. Chng c phn
thn trong sut, thun di, v hu ht c chiu di nh hn 4 cm, thng khng di
chuyn trong nc, nhng c kh nng phng rt nhanh ti con mi. Cc mu kitin
nm xung quanh ming ca chng lm nhim v bt mi - trong c c ng vt ph
du loi nh. Chng khng kn thc n m thng n loi thc n phong ph nht
ngay vng chng sinh sng.
96


Hnh 5-4: Phiu sinh vt Planktonic Crustacea (ng vt thn gip Calanoid)

Thc t ch c mt vi loi trong ngnh thn mm l thuc loi ph du hon
ton. Mt s loi c th thu mnh vo ci v xon c nh (<10 mm); s khc c phn
v nh hn hoc hon ton khng c v v l nhng ng vt tng i trong sut vi
chiu di ln ti 50 cm.
b)- Ph du tng giai on
Mt s ng vt khng xng sng bin khng c giai on u trng bi t
do. Ngay khi n ra t trng c hnh dng ging nh khi trng thnh v sng di
y bin, hoc c trc tip t m. Nhng khong 70% loi sng y ny ra
trng hoc phi trong nc v t n ra u trng, chng l mt phn ca qun th
ph du. Tu tng loi, cc u trng c th mt khong vi pht cho ti vi thng (hoc
thm ch ti vi nm) sng dng ph du trc khi trng thnh nh c y v
bin i thnh dng trng thnh. Trong thi gian ny, u trng tri ni trong dng
nc v c phn tn i xa.
Mt s u trng thng gp ca ng vt khng xng sng y bin nh
Hnh 5-5. c v trai sng y ra u trng c v (loi u trng u bc dim bi)
c mng lng t va bi va bt thc n. ng vt chn t khng cung c cc
giai on u trng bi trong nc (thng l su giai on), tng t nh u trng chn
cho v gip xc ph du khc nhng c phn ra mc pha trc ca b xng ngoi;
cc giai on u trng ny pht trin nh mt b phn gn vi mt cht nn v cc
dng bin thi i ti giai on trng thnh. Sao bin, nhm bin, da chut bin, v
cc loi ng vt da gai sng di y c rt nhiu dng u trng ph du theo tng giai
on, mt s c minh ho Hnh 5.5.
97


Hnh 5.5: u trng ph du tng giai on ca cc ng vt khng xng sng y. (a)
u trng veliger ca c (b) Polychaete trochophorr, (c) u trng ca poWeete; (d)
bipinnaiia ca sao bin); (e) echInopluteus ca mt nhm bin; (f) u trng ca loi
chn t; (g) barnacle cypris: (h) cua zoea; (i) cua megalopa.
ng vt y khng xng sng trong vng nc nng thng c cc giai on
u trng tri ni, nhng cc loi sng y bin su thng khng c giai on ny
m trc tip ra con c dng ging nh khi trng thnh. iu ny c th do vng
nc su thiu thc n ni l lng thch hp cho u trng ph du. vng bin ngoi
khi ni c nc lnh v n ho, u trng ni tng giai on ca cc ng vt y
khng xng sng thng xut hin theo ma ng vi thi gian c nhit cao v
nhiu thc vt ph du. vng nhit i, s sinh sn ca ng vt y khng xng
sng c phn lin tc hn, nhng ma sinh sn r gn lin vi thi gian mi trng c
s bin i, chng hn nh ma; nhng vng nh vy, u trng ph du theo giai on
c th xut hin quanh nm, nhng vi mt khc nhau.
Trng v u trng c (hnh 5.6) cng gp phn quan trng trong qun th ph
du theo giai on; chng c gi l ph du ca c. Mt vi loi c trng vo tng
nn y; chng hn nh c hi thng vi trng xung ct di sui, cn c trch
trng vo to bin hoc ngay di y bin. Tuy nhin, nhiu loi c bin trng tri
t do, nhng loi ny gm c mi, c ng v nhiu loi c c gi tr kinh t cao khc.
Trng tri ni ca cc loi c ny thng c hnh cu, trong sut, v c kch thc nh
vi ng knh khong 1-2 mm. Trng c cha non - ngun dinh dng cho phi
98

pht trin v n u trng. Mt vi loi trng cn cha cc vin du, tr gip cho chng
ni d dng hn.
Cng vi u trng ph du theo giai on ca cc ng vt y khng xng
sng, s xut hin ca trng c trong qun th ph du ph thuc vo chu k trng
v thng gn lin vi iu kin thay i mi trng. T l pht trin ca trng tu
thuc vo tng loi c nhng ch yu l nhit nc bin ni chng sinh sng, nc
lnh s lm qu trnh n trng chm li. Trng thng n trong vng vi ngy ti vi
tun sau khi . S trng c ra c th rt nhiu, v d, mi con ci thuc loi c
bn sao khong 250.000 trng; c fin 500.000, c tuyt 1 triu. Nu em nhn cc
con s ny vi s c , th ta s c mt s trng khng l. Ch ring eo bin nc
Anh, s trng c mi cm c tnh khong 4 x 10
14
. Tt nhin l ch mt phn trong s
ny sng n giai on trng thnh; trng ca nhiu loi l ngun thc n quan trng
cho ng vt ni hon ton v c c ln khc.
V kch thc ca trng, c th phn lm hai loi. Mt s loi trng c kch
thc ln, nhng s trng li t do hn ch v ngun nng lng. Trng to hn cha
nhiu non hn v con n ra cng s to hn so vi trng c kch thc b. Con to hn
thng c t l sng st cao hn, v chng qu to so vi mt s loi n tht ngoi
khi. Mt khc, nhiu loi ra mt lng ln trng cha rt t hoc khng c cht
dinh dng cho phi pht trin. Nhng con c sinh ra t trng ny c kch thc
nh v d b cc con vt khc tn cng. S c con n ra t trng nh b cht nhiu hn
so vi s n ra t trng ln, nhng b li th s trng cng nhiu hn.

Hnh 5-6: Trng c v u trng c ph du tng giai on . (a) trng c trng; (b)
trng c thu; (c) trng c n lng; (d) phi trng c tuyt ang pht trin; (e) u
trng c tuyt mi n; (f) c mi cm mi n.
Khong vi ngy u sau khi n, u trng c vn cn li mt cht non trong
mt ci bao nh di thn v chng tip tc sng da vo ngun dinh dng t bao
non ny cho n khi ming v rut pht trin. Khi non cn kit, u trng bt u
kim n, chng hon ton sng nh vo thc n thch hp l lng trong nc. S ph
99

thuc ny ko di trong sut giai on tri ni ti vi thng, cho n khi chng ln
thnh sinh vt tri ni v c th t kim n mt cch c lp m khng chu tc ng
ca dng nc. ng thi, c con cng d b cc loi khc n tht, tn cng (ng vt
ph du ln hn v sinh vt tri ni). Thng t l cht cao nht trong giai on tri
ni, ch c mt phn nh sng st ti giai on trng thnh. c tnh t l cht giai
on u ny vng nc lnh ln ti 99.999 %.
5.2. cc phng php thu thp mu
Cc phng php thu thp ng vt ph du ph thuc vo kch thc ca
chng. i vi loi ln hn 200 m, thng l ko li c mt tht dy qua ct nc.
Li bt ng vt ph du khc nhau v kch thc, hnh dng v c li, nhng u
c thit k bt cc ng vt tri ni hoc di chuyn kh chm. Li dng n
gin nht c dng hnh nn, ng ming c mt vng kim loi, chp c buc vo
mt ci chai. Loi li ny c chiu di, ng knh v c li khc nhau, n c th
c ko dc, ng hay xin qua vng nc. Loi li ny s lc nc v gi li ng
vt trong qu trnh ko. Cc li phc tp hn c trang b thm b phn c th
ng m cc su mong mun, chng c th c gn vi mt ci khung cho php
cc mu nc cc su khc nhau lt vo qua mt ln ko. Vic phn tch cc mu
thu thp c cho bit chi tit v s phn b ng vt ph du theo phng thng ng
ti vng no . V nhiu loi ph du di c theo phng thng ng theo chu k 24
gi, nn thi gian ly mu cng cn c vo nhng thay i theo su di c.
Vic la chn mt loi li c th ph thuc vo loi sinh vt cn nghin cu v
c im vng nc mun ly mu. V d, loi li c mt nh (khong 100-200 m)
ch dng thu thp cc loi ph du cc nh. Tuy nhin, n li khng thch hp thu
cc loi bi nhanh c kch thc ln hn nh u trng c, v chng s mc vo li v
phi c ly ra t t, nu khng li s b rch. vng nc su, ni ng vt ph du
c kch thc ln hn v c s lng t hn, thng dng loi li c mt rt th. Tt
c loi li ny c th c gn mt lu tc k tnh lng nc chy qua trong mt
ln ko; iu ny cho php xc nh s lng ng vt ph du thu thp c.
Thc t khng c mt loi li no c th bt c tt c cc loi ng vt ph
du. ng vt ph du nh hn 200 m (nh v siu nh) khng th ly mu c bng
li; thay vo , ngi ta thu mt mu nc bng nhng ci chai hoc bm ln t
su cn thit v ng vt ph du c lc bng nhng ci lc, my ly tm hay lng
xung. i vi ngnh ng vt nguyn sinh ph du v thc vt ni ta cng c th thc
hin theo cch ny.
Hin nay ngi ta nhn ra rng mt vi ng vt ph du c kh nng trnh khi
b mc li, nh chng nhn thy hoc cm nhn c s xo ng ca nc do li
chuyn ng gy nn. Thm vo , loi ph du dng jelatin cng yu t cho nn
khng th thu c chng mt cc nguyn vn bng li; mt vi loi phn hy rt
nhanh trong mi trng c cht bo qun. Gip xc thng c nhiu loi ph du b
mc vo li v hu ht l chng rt nh nn khng th t bit cch trnh, v v b
100

xng ngoi cng bo v chng khi b cht hay bin dng trong li v cht bo
qun. Do vy m s gip xc c lin quan n cc loi ng vt ph du thng b tnh
qu ln, v ph du thc s mc vo li c th khng cho ta mt con s ng v qun
th ph du nhiu vng. nhiu vng, s ng gp v sinh khi v s lng ca gip
xc, c bit l loi chn cho, cn phi c nh gi li.
Trong nhng nm gn y, c nhiu c gng trong vic quan st loi ng
vt ph du trc tip ngoi thc a, hoc s dng bnh kh p ln su xung c 30
m, hoc tu ngm ni vng nc su hn. Cc cng ngh ny gip cho ngi ta pht
hin ra c nhiu loi mi, c bit l loi yu t, d b tc ng; trnh c vn
nh gi thp s lng v sinh khi ca cc ng vt qua thu thp bng li. Phng
php s dng m thanh sinh hc pht trin t vic s dng thit b pht hin tu in
ngm nh v n c, cng c p dng xc nh v c tnh mt ca ng
vt ph du c ln, nh tm vy - loi sinh sng kh dy c trong nc.
5.3 phn b theo ng vt ph du theo chiu su
5.3.1 Phn b theo chiu su
Chng ta phn loi ng vt ph du theo kch thc, mi trng sng, loi
ph du v thi gian sng ph du. Chng cn c phn nhm theo khu vc sng
trong ct nc.
Cc loi sng vnh vin trn mt nc bin v c th chng l ra mt phn
trong khng kh c gi l pleuston. i khi chng c xem nh mt loi c bit v
chng b gi y i ch khng phi l nc. Neuston bao gm cc loi sng su
ch vi chc milimet trn mt nc. V mt sinh thi hc, kh c th tch bch mt
cch r rng cc loi ny, v y, ta ch tho lun mt cch tng th v cc sinh vt
sng b mt i dng. Qun th ny pht trin mnh vng bin nhit i, v hu
ht cc v d sau y u l cc loi sng vng nc m.
V d v pleustonic: chng bao gm sa c lng chm Physalia sng theo qun
th, v Velella cng h hng ca chng, tt c u c bong bng kh nh ln khi mt
nc. Cc xc tu di lng thng ca Physalia gip chng bt sinh vt ni v c con
di mt bin rt tt. Velella c cc xc tu ngn v bt thc n (chn cho, u trng
ca cc loi gip xc, trng c v trng tm vy) rt gn mt nc. Mc d c xc tu
chm nga, ra bin v cc loi nhuyn th sng st mt u n loi sa c lng chm.
Janthina l mt loi c to ra mt mng bong bng khng kh bc trong lp nhy; n
treo ngc mnh trn gn mt nc v n n c Physalia, Velefla cng cc ng vt
sng gn b mt khc. Glaucus - mt loi c sn bin treo ngc mnh st mt nc
bng vic thu khng kh sau gi trong nhng ci ti c bit ca ng tiu ho. N
cng n Physalia v Velefla v tiu ho c t bo chm nga v cc m khc. c sn
bin c kh nng hp th t bo chm nga rt tt m khng h phi thi ra ngoi, cc
t bo chm nga sau c cc lng t chuyn ti nhng ci ti c bit ngay u
nhng phn nh ln t lng ca c. Nhng ngi i tm bin chm phi loi Glaucus
101

ny ni rng h b au v nga ko di n vi ting ng h. Mt s chi khc ca
cng ng neuston bao gm mt s cn trng thng thy vng bin khi; chng
thuc vo b cn trng Hemiptera v loi Halobates. Nhng loi cn trng khng cnh
sng trong nc ny khng th sng ngp chm trong nc v do ch n cc loi
sinh vt ngay b mt, bao gm enidarians v neustonic: cc loi chn cho.
Mt s neuston, c bit l u trng cua v c, rt trong v do kh c th
phn bit bng mt. Nhng nhiu ng vt neuston nhit i (chng hn nh Velella,
Halobates, Janthina, Glaucus) c mu sc rt ni bt nh mu tm hay mu xanh
dng. Ngi ta cho rng nhng mu ny gip cho chng trnh c nh hng ca
tia cc tm mc mnh, nhng cc mu ny cng c th dng ngu trang rt
hiu qu nhm trnh b n tht v chng ho ln vi mu xanh ca nc bin vng nhit
i. Janthina v Glaucus cng c b ngoi tng phn nhau; tc l, b mt phn hng
xung nc c mu sng hn so vi phn gn mt nc, v cng c th l chng c
mu sc nh th t bo v mnh trnh khi b n tht. Nhng sinh vt n tht bin
nh c nhn vo cc ng vt thn mm ny t di ln v mu sng phn di ca
loi thn mm ln vi mu nn sng ca tri, trong khi cc sinh vt n tht trong khng
trung, th mu xanh sm hn b mt trn ca con mi li ho ln vi mu xanh m
ca nc bin. Mu sc thay i khng phi l cch bo v duy nht ca neuston.
Chng hn nh, mt s loi chn cho neuston c kh nng nhy ln cao khi mt
nc trnh b n tht.
Mt qun th sng mt nc c bit pht trin bin Sargasso. Sargassum,
mt loi to bin, to nn mt mi trng sng thun li cho tp hp duy nht c hn
50 loi ng vt. Tng khi lng to Sargassum ti y c c tnh khong 4
n 10 triu tn. Nhiu loi sng y ch yu l ng vt y gm thu tc, chn
ngng bin, cua, tm v cc loi gip xc khc. Mt vi loi c hu (ch sng ni c
mi trng c bit), bao gm mt s loi cua, tm pht trin kh nng ngu trang
bo v bng cch bin mnh v hnh dng v mu sc ging vi to Sargassum hn.
Vng nm ngay di mt nc su ti 200 n 300 m c coi l vng st mt
(epipelagic). Nhiu loi ng vt ph du l c dn c nh, mt vi loi khc di c ti
vng ny vo ban m. Ch c nhng loi ph du sng su nh hn 300 m vo
ban ngy mi c coi l ph du gn mt thc s. Vng st mt trng vi vng cc
vng euphotic v disphotic, to iu kin thun li cho s sinh si ny n v a dng
sinh hc. Nhiu loi n thc vt v n tp sng vng ny, bao gm cc loi gip xc
nh (nh chn cho), v u trng phu du cc nh. Nhiu loi trong s ny c dng
trong sut.
Vng mesopelagic nm gia vng gn mt epipelagic v su khong 1000
m, v nhng loi sng y vo ban ngy c gi l loi mesopelagic. Mt vi loi
sng vng ny nh tm vy, t nht c mt phn l n thc vt, v chng chuyn ln
vng cpipelagic vo ban m n thc vt ni. Tuy nhin, nhiu loi khc ch n tht
hay n cc cht mn phn hu, chuyn tm n cc ht thc n c kch thc ln hn.
Cc cht hu c dng ht ri t trn xung, c bit l cc cn lng, thng tch li
102

su 400 n 800 m nh gradient mt . Ngun thc n phong ph ny s b phn
hu bi vi khun, nhng ng thi y cng l thc n ca cc loi ng vt ph du
khi chng pht hin ra v dn v cc vng ny sinh sng, v s phn hu cng vi
qu trnh h hp ca ng vt s khin cho oxy b hao ht nhiu. Cc hot ng sinh
hc gp phn hnh thnh cc tng c oxy thp, hoc cc vng xy c th gim t
mc bnh thng l 4-6 mg/l xung t hn 2 mg/l. Cc yu t vt l cng lin quan n
s hnh thnh cc tng c xy thp ny. xy c b sung trn mt bin qua s
tip xc vi kh quyn, v c chuyn xung vng su do nc trn mt cha y xy
s chm xung di. Nh vy, tng c nng xy nh nht biu th lp nc c s b
sung ngun xy nh nht, cng nh vng c s h hp ca ng vt ln nht.
C xu hng l cc loi sng vng mesopelagic thng ln hn h hng ca
chng sng vng epipelagic, v nhiu loi ng vt c v thng c mu hoc
en. V d, tt c cc loi tm ngoi khi sng su di 500-700 m vo ban
ngy th u c mu ti, trong khi cc loi sng nc nh hn th trong sut
hoc gn nh trong sut. Nhiu ng vt ph du mesopelagic (v c) cng c mt to v
nhy cm vi nh sng c bc sng xanh lc-xanh dng; y l cc bc sng t
bc x xuyn c su nht v cng l quang ph ca hu ht cc nh sng pht quang
sinh hc.

Hnh 5-7: Sinh khi ca ng vt ph du c bt bng li (khng bao gm loi sa
c lng chn v salps) k t mt nc cho ti su 8000 cm. vng ty bc Thi
Bnh Dng ((45
O
N) (ng m) v vng nhit i Thi Bnh dng.
H thng phn loi theo su da trn c s cc iu kin mi trng thay i
theo chiu su ca bin, v ng vt sng su khc nhau c cch sng khc nhau.
Tuy vy, c nhiu ng vt di chuyn t vng ny sang vng khc, v nhng vng su
hn thng kh c th c phn nh mt cc rch ri. Ngoi ra, s phn b cc loi
103

theo su c th thay i theo v . iu ny c bit ng vi nhng loi sng
vng nc lnh c kh nng thch nghi cao vi s thay i p lc thu tnh. Chng hn
nh loi hm t Eukrohnia hamata, sng ngay st mt vng cc, nhng nhng
vng gn xch o hn th chng li sng di nc su v lnh.
5.3.2 S di c theo chiu thng ng
Mt trong cc c im phn ng ca sinh vt ph du l di c theo chiu thng
ng vi chu k 24 gi. Di c theo chiu thng ng l hin tng cc sinh vt di c
ln mt thong vo ban m v chuyn su xung vo ban ngy. iu ny c bit
n t thi Challenger Expedition, nhng cho n nay chng ta vn cha hon ton
tho mn vi nhng gii thch v hin tng ph bin ny v tm quan trng v mt
sinh thi ca s di c theo chu k 24 ting . Tt c cc nhm ng vt ph du (gm
c cc loi nc ngt), u c t nht mt vi loi di c theo phng thng ng, chng
hn nh trng roi v nhiu loi sinh vt bi, bao gm c ng vt chn u v c.
Nh s di c ny, khi so snh gia mu thu c ca sinh vt ph du ngy v
m c thc hin cng mt vng v cng su, ngi ta s thy c s khc
nhau v cu to ca tng loi v tng sinh khi ca chng. Chng hn nh, ta c th
thy s khc nhau hnh 5.8.
Mi loi c khong cch di c vo ngy v ban m ring, v khong cch
ph thuc vo tng giai on sinh trng v gii tnh ca tng loi (v d nh loi
Calanusfinmarchicus con ci trng thnh di c rt xa, trong khi con c th km
hn nhiu). Khong cch di chuyn cng thay i theo ma, v tr a l v iu kin
thi tit (chng hn, nh my, bo t, v.v...). Tuy vy, nhn chung c 3 dng di c ca
ng vt ph du bin l:
1. Di c ban m: i ln mt ln trong ngy, thng l bt u vo lc mt tri ln v
i xung t cc tng pha trn, gn lc mt tri mc. y l kiu di c thng dng
nht c 101 loi ng vt ph du bin.
2. Di c lc nh sng m t c hai ln i ln v hai ln i xung. C mt ln i ln
lc mt tri ln ti mt su nh nht vo ban m, nhng li cng c mt ln i
xung vo ban m. Khi mt tri ln, cc ng vt li chuyn ln gn b mt v sau
li i xung ti su ban ngy.
3. Di c o ngc l kiu di c t gp nht. Kiu di c ny c mt ln i ln mt
nc vo ban m v mt ln i xung vo ban ngy ti su ln nht.
Di c theo chiu thng ng chnh l nguyn nhn sinh ra cc tng phn tn
di nc su. Cc tng ny c th chuyn ng sut 24 gi, v nhp hot ng ca
chng cho thy nguyn nhn chnh l s chuyn ng ca ng vt. Vo ban ngy c
khi ln ti 5 tng phn tn su khong 100 - 750 m. Ban m, cc lp ny chuyn
ln mt nc v phn tn ra, hoc chng nhm li thnh mt di rng tri xung n
150 m. Cc lp phn tn theo su ny c v do s chuyn ng ca cc loi gip
xc c kch thc ln (v d nh tm vy, tm thng) v c c bong bng cha kh c
104

kh nng phn m, nhng cc loi ng vt ph du khc nh cc loi chn cho ln
him khi to ra c nhng lp phn m nh th.
Thc t hin tng di c theo phng ng chu nh hng ca nh sng t
nhin: theo chu k bng ti cho thy thay i cng nh sng ng vai tr nh mt
tc nhn kch thch s di c. Cng nh sng thay i t nhin theo ma hoc thm
ch trong ngy c th lm thay i su m mt loi no sinh sng. Di nh sng
lin tc vo ma h Bc cc, s di c c th hon ton khng c. Trong trng hp
nht thc s khin cho ng vt di c ln trn vo ban ngy khi cng nh sng
gim. trong phng th nghim, thi gian di c c th thay i hoc khng ty theo
nhng th nghim thay i chu k sng v ti. Ngoi nh sng, cc yu t khc cng
c vai tr trong vic to nn hin tng di c trong ngy; v d do i, ng vt s di
chuyn v vng c ngun thc n di do hn di s bo v ca bng ti.
Nhiu gi thuyt c a ra tr li v s di c ny, nhng khng thc t
khi gii thch c cu di c ca mi loi theo phng thng ng mc ton cu.
iu quan trng l phi bit rng cc gi thuyt di y c th khng ng cho tt c
m mi gi thuyt c th ph hp cho mt s loi ny hn cc loi khc.
1. Mt gi thuyt cho rng ng vt sng trong bng ti hoc gn nh ti sut 24 gi
t b nguy him do cc loi n tht sng trong nh sng. Chnh v vy m c s di c
i xung vo ban ngy. Cn di chuyn ln pha trn l gip chng kim c nhiu
thc n hn. Tuy nhin, nhiu ng vt c su di c vo ban ngy nm trong ct
nc c chiu sng, nhiu loi khc th pht quang sinh hc v do vy m chng
c th b pht hin vo ban m.
2. kin th hai cho rng ng vt ph du bo ton nng lng bng cch khi khng
n th xung ng vng nc su v lnh hn, ni nhu cu nng lng cn cho
trao i cht t hn. Ngi ta cha chng minh c rng nng lng tit kim
c trong mi trng lnh c th b p li nng lng s dng bi trong qu
trnh di chuyn. Tuy nhin, nng lng cn thit bi l rt t, thng ch vi phn
trm ca nng lng trao i cht c bn.
3. Gi thuyt th ba ni rng, ng vt ph du di c theo phng ng l chu tc
ng bi cc lung nc chuyn ng theo nhiu phng khc nhau vi tc
khc nhau. Do vy, mi mt ln di chuyn ln l chng vo mt mi trng kim
n hon ton mi. Vng kim n mi c th thc n nhiu hn hay t hn vng m
chng sng vo ban m, nhng nh s di chuyn theo phng ng m cc sinh
vt nh c kh nng linh ng km c th trnh khi phi trong mt vng c
ngun thc n khan him cng nh trnh khng b tp trung qu ng ni c
ngun thc n di do. Tuy nhin, nhiu th nghim thc hin bng cch th thc
n vo cc vng vi nng khc nhau a li nhiu kt qu mu thun, khc
nhau i vi tng loi. mt vi trng hp, ngun thc n km khin cho hin
tng di c theo phng ng bin mt; nhng cng c trng hp ngc li.
105

Cho d gi thit no i na, s di c theo phng ng trong ngy c mt s h
qu rt quan trng v mt sinh thi hc. Mt l, do tt c cc c th ca mt loi khng
di c ng lot vo cng mt thi im v ti cng mt su, mt qun th s mt i
mt s c th v cng c th thm mt vi c th khc. S pha trn cc c th t cc
qun th khc nhau tng cng s pha trn v gen v iu ny c bit quan trng i
vi cc loi c kh nng km di c theo phng ngang.
Mt kt qu quan trng khc ca di c theo phng ng lm thc y s
chuyn ho ca cc cht hu c sinh ra trong vng st mt ti vng su hn. Mi mt
loi di c n thc n khong nc nng vo ban m v n ban ngy lng thc n
ny c chuyn ti vng su hn. Loi n c n thc vt ph du vng st mt, sau
chuyn ti cc vng su hn v chng thi cc cht mn hu c vo , v y
cng l ni chng c th lm mi cho cc loi n tht vng nc su. Loi n tht v
loi n cht thi ra ti lt mnh li thc hin nhng chuyn di c theo phng thng
ng ti cc su ln hn. Qu trnh vn chuyn cc cht hu c theo phng ng
ch yu do bn thn ng vt chuyn, hoc do cht thi khc vn chuyn nhanh hn
nhiu so vi qu trnh chm t nhin.
5.4 thay i kiu theo thi gian
Cc kiu bin thi theo ma ca ng vt ph du c lin quan ti s bin
ng ca thc vt ph du, mt vi loi di c thng ng thay i theo ma v c lin
quan ti chu k sinh v su thch ng vi tng giai on trong chu k sinh
trng. bc Thi Bnh Dng, cc loi chn cho thng gp, c s thay i rt r v
su. ngoi khi vng bin Canada, Neocalanus plumchrus trng thnh v
tri qua ma ng su 300-450 m, v trng vo khong thng 12 ti thng
4. Trng ni ln b mt, u trng n v trng thnh cc su trung gian. u trng
c mt vng nc gn b mt t thng 2 n thng 4, v chng pht trin ln giai
on V t thng 3 ti thng 6, khi s sinh sn t mc cao nht. Vo u thng 6, cc
c th giai on V cha mt khi lng ln lipit do tch lu dn khi n thc vt ni,
v chng bt u di c ti vng nc su hn ni chng s s dng dn ngun nng
lng c tch tr t trc. chng pht trin ln giai on VI, giao phi, v
trng trong ma ng. ngoi khi, chu k sinh trng thay i c khc mt cht,
chng trng vng nc su (>250 m) t thng 7 ti thng 2 v cc giai on u
trng u bt u xut hin vng nc trn 100 m vo thng 10. Tuy nhin, n tip
tc di c theo ma gia vng nc gn b mt, ni u trng pht trin, v vng nc
su ni chng bt u giao phi v trng. Mt kiu di c theo phng thng ng
tng t c lin quan ti cc giai on sinh sn khc xy ra loi Neocalanus, mt
loi chn cho ln cng kh ph bin bc Thi Bnh Dng: con trng thnh sng
vng nc t 1000 ti 2000 m, v trng ni su hn; cc giai on u chng
thng vng trn su 250 m.
106


Hnh 5-8: Thay i theo ma vo ban ngy (trng) v ban m (en) theo phng
thng ng giai on V v VI ca hai loi Cakimis helgolandicus (a) v C
finmarchicus (b), vng bin Celtic. S lng mi m ko li cc su 5 m, c
thc hin vo ban ngy.
Hnh 5-8 th hin c di c thng ng theo ngy v theo ma hai loi chn
cho n thc vt vng bc i Ty Dng, Calanus helgolandicus v C.
finmarchicus. V ma ng bin Celtic, u trng chn cho giai on V v VI c
hai loi phn b kh ng u t mt nc cho ti su 100 m, v c s khc nhau
rt nh gia ban ngy v ban m. Vo ma xun (thng 4), c hai loi bt u tp
trung vng nc cn. y chng thc hin s di c theo phng thng ng theo
chu k 24 ting. Vo thng 7 v thng 8, ng phn nhit hnh thnh r v hai loi c
s khc nhau rt r trong phn b. Calanus helgolandicus tip tc pht trin trong
vng b mt nc m v vn di c theo ngy, nhng C. finmarchicus di chuyn xung
vng nc su lnh hn di ng phn nhit v lc ny c s khc nhau trong
su m chng xung gia ban ngy v ban m. Vo cui thng 9, c hai loi sng
vng nc su hn 40 m vo ban ngy v C. helgolandicus tip tc di chuyn theo
phng thng ng ln mt nc vo ban m.
Tm Nam cc (tm vy) cng di chuyn theo su trong chu k sinh trng
ca chng. Trng tm t li trn b mt nhng chm rt nhanh xung su 500 -
2000 m v n . Sau u trng tm t t ni v bi ln b mt v s pht trin
hon ton s xy ra y, tm con v tm trng thnh sng ngay b mt hoc
107

st mt. Ton b chu k sng ca chng khong t nht 2 n 4 nm. Trong thi gian
ny, s di c theo phng thng ng cc giai on khc nhau v dng i dng
chy theo nhiu hng a chng i ra v ti Nam cc. Mt vi loi chn cho v hm
t Nam cc thng thy cng di c thng ng theo ma tng t: vo ma h cc
nam, cc loi ny xut hin trn mt nc chy v hng bc t vng Nam cc; vo
ma ng, li thy chng trong dng i dng su hn chy v hng ngc li.
S di c mnh m theo ma thng xy ra cc loi sng vng nc lnh v
n i, hoc vng nc tri. Di c gip cho cc ng vt ph du giai on cha
pht trin ti c vng b mt chng c th nhn thc n v pht trin. vng
nc n i, do s sinh sn trn b mt gim vo ma h v ma thu, nn u trng
cc giai on cui hoc trng thnh ang thi k sinh sn chuyn xung vng
nc su hn. y, trong vng nc lnh v khng c iu kin sinh sn chng c
th bt u mt trng thi gi l diapause, trong , s trao i cht din ra rt chm
v chng khng n. Thay vo , chng tn ti nh ngun nng lng d tr trong thi
gian chng vng mt.
C di c trong ngy v di c theo ma a cc sinh vt vo dng chy i
dng theo cc hng khc nhau vi tc khc nhau. D vy, cc qun th cc ng
vt ph du bin vn nhng vng c c tnh ph hp vi chng. Mt kiu khc ca
di c thng ng theo ma mt loi c th l bo m s duy tr trong mt mi
trng thch hp, hoc trong vng c iu kin sinh sn (nh mt s trng hp i
vi mt vi loi c lin quan n chu k sinh sn hng nm ngoi khi pha ty ca
bc M v chu Phi). Tng t di c trong ngy c th gi cho loi ph du li mi
trng sng thch hp. V d, di c trong ngy ca cc loi vng ca sng l c
th thch nghi vi dng triu ln xung v tr li trong vng ca sng. Trong mi
trng hp, s chn lc t nhin thng gi li nhng c th di c vi thi gian thch
hp; bt k c th no vi kiu phn ng khng hp vi h thng t nhin s c xu
hng bin mt. Kt qu tt yu ca iu ny l t nht mt s loi th hin kiu di c
thng ng khc nhau cc vng khc nhau, chng c th thay i m bo s tn
ti ca qun th theo cc vng a l. Nhng kiu di c khc nhau cc loi c th l
s thch nghi tin ho duy tr ni ging trong iu kin mi trng thch hp cc
ch dng chy khc nhau.
5.5 cu hi v tho lun

108

Chng 6
sinh vt Tri ni
6.1 Phn loi
6.1.1 Khi nim v sinh vt tri ni (nekton)
Sinh vt tri ni l nhng ng vt c th bi vt c sc chy ca dng i
dng. Sinh vt tri ni thng sng vng c cc iu kin tng ng vi s Reynold
cao. Sinh vt tri ni bao gm cc loi c, ng vt thn mm, ng vt c v b
bin, chim v b st. Khng ging nh sinh vt ph du, chng c th bi, v chng li
c sc cun ca dng chy mnh. iu ny cho php chng di chuyn vi khong
cch kh xa trong vng mt ngy. Mt s sinh vt tri ni di c ti hn hng ngn km.
Sinh vt tri ni c ng lng trong khi di chuyn di nc. Hay ni cch
khc, nu mt con c ang lao nhanh di ln nc m ngng li, th n s khng
dng ngay lp tc m nhp nh mt lc. Khng c mt lp ranh gii tng i dy no
bao quanh n tng tc vi mi trng cht lng xung quanh. Thc t ny cho
php sinh vt tri ni lao rt nhanh. iu ny ngc vi ph du, b cn bi nht ca
cht lng. Khi mt sinh vt ph du di chuyn trong nc, n s ko theo mt lp nc
mng bao quanh.
Ngc vi sinh vt ph du, sinh vt tri ni c kh nng di chuyn rt tt,
chng c th cng li sc cun ca dng i dng hay cc dao ng ca nc do
gi sinh ra. Chng hu ht u l cc ng vt c kch thc ln gm c cc loi ln
nht v di chuyn nhanh nht bin. ng vt ph du ch yu l cc ng vt khng
xng sng, cn sinh vt tri ni ch yu l ng vt c xng sng. C l sinh vt tri
ni ng o nht tnh c theo s loi cng nh s c th, nhng ni chung tt c cc
lp c xng sng tr loi lng c u c i din l sinh vt tri ni. c trng ni
bt ca chng l kh nng bi vt qua c dng nc m khng ph thuc vo mi
trng sng. Trong chng ny, chng ta s ch xem xt cc ng vt tri ni sng
vng gn mt nc ngoi khi, gi l sinh vt tri ni bin khi (oceanic nekton).
Sinh thi hc v s thch nghi ca sinh vt tri ni sng vng gn b, chng hn nh
lp to b y v rn san h v sinh vt tri ni vng nc su s c ni ti trong
cc chng tip theo. S thch nghi ca sinh vt tri ni bin khi rt c bit, khc vi
s thch nghi ca sinh vt tri ni sng vng nc su v gn b, do vy cn nghin
cu ring.
Nhng ng vt ny bi nhanh v c mt tt c cc i dng, nn vic
nghin cu chng ngay trn bin l rt kh v khng th gi chng li trong mi
trng t nhin c. Ngi ta bit rt t v cc kha cnh sinh thi hc hay lch s i
sng ca cc sinh vt . Do thiu cc ti liu v khng c nhiu kt qu th nghim,
109

nn phi phng on sinh thi hc ca chng bng cch nghin cu cc c im sinh
l v cu trc gii phu cc sinh vt mu.
Cc th d in hnh v sinh vt tri ni c cp di y (xem hnh v 6.1):
ng vt chn u thuc ngnh nhuyn th, thng l loi n tht, c c im
phn ng phc tp v mt h thn kinh c cu trc kh cht ch, ng thi c
nhng cnh tay bt mi v mt ci ming rt khe. Mt s ng vt chn
u c th iu khin sc ni ca chng bng cch thay i lng kh cha
trong mt ti rt chc chn.
C bin l loi c mt hu ht cc mi trng sng bin, ch yu l c c
sn (c mp, c ui) v c c xng.
ng vt c v bin bao gm c voi, hi cu, s t bin,...
B st bin vi cc loi ni ting nh ra bin, rn bin...

(c)
(d)
(e)
Mng bin
lng en
C voi xm
B st bin
kemp's ridley

Hnh 6-1: Trong hnh ny (a)- mc Rossia pacifica (thuc ng vt chn u), (b)- C
n tht nm gi cht (c), (c)- mng bin ln lng en (chim bin), (d)- c voi xm
(ng vt c v bin), v(e)- kemp's ridley (b st bin).
Cc yu t mi trng vng gn b mt tc ng ti sinh vt tri ni sng
y cng ging nh ni ti chng trc v sinh vt ph du, bao gm nh sng,
nhit , khi lng ring v dng i dng. Tuy nhin, tm quan trng ca yu t
i vi s thch nghi v cch sng ca sinh vt tri ni c khc mi nhau. Cn ch
rng, mi trng i vi mt loi c ln v bi nhanh hay ng vt c v l khc i
vi loi chn cho nh.
Mt s iu kin mi trng c vai tr kh quan trng i vi sinh vt tri ni
v s thch nghi ca chng. Trc ht, y l mt vng khng gian ba chiu rt su v
110

rng. Th hai, y khng c lp nn cng v ln, do vy ng vt lun ni trong mt
mi trng trong sut khng c s bo v c th trnh khi b n tht. Chng khng
c ni khi di chuyn t ni ny sang ni khc theo phng ngang. Cui cng,
thiu lp nn cng c ngha l khng c mt tm chc chn cho ng vt, nn hu
ht nhng loi ny c cu trc tht chc nng hn nc bin xung quanh.
Khng gian ba chiu trong iu kin khng c chng ngi g l iu kin
thun li cho ng vt c kh nng thch nghi nhanh chng. Kh nng di ng cao v
vt c cc qung ng xa gip cho ng vt la chn s pht trin ti u cho h
thn kinh v cm gic cung cp v iu khin thng tin cn thit trong khi di ng,
tm kim thc n v trnh cc mi nguy him.
Tng t nh vy, trong iu kin thiu ni n nu v c kch thc ln chng
pht trin kh nng bi nhanh trnh b n tht, ng thi cng tm thc n.
Thiu ni n np cng lm pht trin kh nng ngu trang ca sinh vt tri ni. c
th gi mnh c t trng thn th ln trong mi trng nc c t trng nh hn,
chng phi pht trin mt s kh nng thch nghi c th ni c trong nc.
Kh nng bi ca c vng gn b mt gip cho chng trnh c cc dng i
dng v chng c th di c t vng ny sang vng khc, la chn c ni c mi
trng sng thch hp, l ni c ngun thc n di do hay c iu kin thun li
cho sinh sn. Nhiu loi c th di c t hng trm ti hng nghn km t ni kim n v
ni , mt s loi khc li c th di c gia vng nc mn v nc ngt. Chng hn,
c Anadromous, nh c hi, c tm, c trch, c t-me v c mt trng vng
nc ngt. C con sau di c ra bin, v sng gn nh sut cuc i ca chng .
Thi gian sng bin tu thuc vo tng loi, nhng khi trng thnh, chng li
quay v vng nc ngt trng. Mt s loi nh c hi Thi Bnh Dng, cht
sau khi giao phi; nhng cc loi khc, nh c hi i Ty Dng, th khng nh vy,
chng c th quay tr li ni vi ln.
6.1.2 Gip xc (Crustacea)
Mc d c mt s loi tm v cua bi vng gn b thuc vo loi sinh vt tri
ni, nhng c im sinh hc v cc loi khc nhau vn cha c bit nhiu, v rt t
loi trong s cc loi tm cua ny c kh nng sinh sn cao c th tr thnh ngun
ti nguyn c th khai thc. Chn mi lm phn trm cc loi gip xc c th khai
thc thuc loi sinh vt y. Tuy nhin, ngi ta ang ch hn n cc loi tm vy,
v nhng loi ny pht trin rt mnh.
Vi s suy gim s lng c voi v ngh sn c voi, Euphausia superba, mt
loi tm Nam cc - ngun thc n ch yu cho c voi sng hm, tr thnh mc
tiu ca ngnh hi sn. Nhng con tm vy ln ny (di 5-6 cm) thng c sy kh
v ch bin nui gia sc, gia cm v c trang tri. Tm bt u c nh bt
Nga v Nht vo nhng nm 60 v ngi ta thy c s thay i rt ng k v s lng
nh bt vo nhng nm sau . Nm 1986, ngi ta bt c 446.000 tn, mt con s
rt khim tn nu so vi lng tm b cc loi n tht t nhin tiu th (c tnh hng
111

nm c khong 470 triu tn). Lng tm c th nh bt c c tnh t nht l 25-
30 triu tn mt nm, hay ni cch khc l bng mt phn ba tng lng c c nh
bt trn ton th gii hin nay. Chi ph nh bt Nam cc kh cao, mc d tm
sng thnh tng n ln, nhng cng phn tn ra kh rng v i khi su 150 -
200 m. Tuy nhin, mt khi nh v c n tm nh phng php phn m, th ch
cn mt m vt bng mt ci thng chuyn dng ln l c th thu c 10 tn tm.
nh hng v mt sinh thi hc t vic nh bt lng ln tm n cc qun th h
sinh thi Nam cc (k c s phc hi ca c voi) th cha r nhng v tm nm trung
tm ca li thc n Nam cc, nn cn phi thn trng khi m rng nh bt.
Mt loi tm vy khc cng c nh bt vi mc ch kinh t l Euphausia
pacifica (Tm vy thi bnh dng), dc b bin Nht Bn. nh bt tm y rt
khc v vo ma xun, tm sng thnh n trn mt nc, do vy rt d bt. Tm sau
c ch bin lm thc n cho nui c. Tm vy l ngun protein v vitamin A rt
di do; ngi ta cho rng chnh vitamin A gip tng cng cu trc v sc t cho tht
ca c nui.
6.1.3 ng vt chn u (Cephalopoda)
Nh ni trn, ng vt chn u thuc ngnh thn mm. Chng l loi thn
mm c kh nng di chuyn tt nht, c s iu khin ca h thn kinh vi t chc kh
cht ch, chng l loi n tht. Mc c th tn cng con mi bng cch co bp cc c
bn ngoi v tng nc qua ng. ng mc phi c hng nh th no y n c
th tn cng bng cc ru mc pha trc. Sau , n cn con mi bng ming. Con
mi c th b tm bng ru, mc ng v loi tm ru, ru ca chng c nhng hng
gic. Mc cng s dng hai xc tu di a con mi vo ming. Bch tuc th khng
c gic nhng c hn 90 ci ru, trong khi mc c 10 chic (8 ngn v 2 di). Loi
mc Sepia thng n cc loi gip xc y bng cch pht nc vo ct pht hin ra
con mi v khi con mi di chuyn, chng s b tn cng.
ng vt chn u c kh nng i mu rt tt. Kt hp ba mu c bn, mc
thay i mu rt nhanh, m iu ny thng lin quan n mc ch ngu trang trnh
b sinh vt n tht pht hin. S thay i ng ch nht l kiu vy sng ca b mt
ngoi ca thn. Ngi ta cho rng b phn ny nhm lm cho n ging vi cc con
sng ln tn gy ra bi nh sng chiu xung y bin.
ng vt chn u sinh sn do s chuyn i b sinh tinh t con c sang con
ci v s th tinh bn trong. hu ht cc loi, trng c trn mt lp nn hoc
tri ni t do. Mc giy ci (Argonauta argo) vn hai chic ru v tit ra mt chic v
hnh xon c rt p giu trng trong.
Mc, mc ng v bch tuc l cc chi thuc ngnh thn mm, to nn lp ng
vt chn u. Mc chim ti 70% lng ng vt chn u nh bt c, v lng
ny ang tng mt cch ng k. c tnh hng nm c th nh bt c 10 triu tn
mc. Tuy vy, ngi ta vn khng bit nhiu v cc c im sinh hc v sinh thi hc
ca chng.
112


Hnh 6-2: Mt s ng vt chn u: (a) mc Rossia pacifica, (b) mc Octopus
dofleini vng Thi Bnh Dng, (c) Mc ng Sepiella japonica vng ty Thi Bnh Dng
Nhiu loi mc ph bin v ang l mc tiu nh bt v l ngun thc n
chnh cho con ngi nhiu nc trn th gii. Nm 1981, Nht Bn bt u s dng
li tri nh bt mc vng bin Thi Bnh Dng. Li tri l cc tm li kt
bng si n rng 8-10 m v di ti 50 km, kch thc li thng l 90-120 mm.
Vo ban m, ngi ta t li thng ng vng bin khi v cho n tri theo gi
v dng i dng trong khong 8 ting bt c v mc. Cho ti nm 1989, Nht
Bn, Hn Quc, i Loan s dng n 800 thng li tri Thi Bnh Dng
bt 300.000 tn mc mi nm v c khong 200 thng li khc c s dng i
Ty Dng v n Dng. Ngoi mc, nhng chic li gn nh trong sut ny cn
bt mt lng ln cc loi khc. bc Thi Bnh Dng, c hi c ngi ta nh
bt nhiu nht (tri php); nam Thi Bnh Dng, c ng l ngun thu sn c gi tr,
lng nh bt nm 1988 ln n 60.000 tn. Ti nm 1989, ngi ta quan tm hn
n s lng ng vt b nh bt bng phng php ny. Ngoi vic nh bt cc
loi c (k c c mp) mt cch khng c chn lc, ch ring Thi Bnh Dng c
tnh hng nm c ti 750.000 n 1.000.000 chim bin v 20.000 ti 40.000 ng vt
c v bin b git do s dng li.
C th nh mc bng cc phng php c chn lc hn nh Nht Bn, nh
bt hng trm nghn tn mc Todarodes pacificus mi nm bng k thut ch bt
ring mc. Loi mc c bit ny thc hin mt cuc di c ngon mc di ti 4.000
km, t ni chng c n ra bc ca bin ng (khong 32
o
v bc) ti vng ph
cn ca qun o Kurile (45
o
v bc) v li quay v.
6.1.4 B st (Reptiles)
C kh t s loi b st sng bin. c bit n nhiu nht l 5 loi ra bin,
s loi rn bin nhiu gp 10 ln ra v c mt loi thn ln bin, mt loi k n c
bin qun o Galapagos. Mt vi loi c su sng gn b, loi ln nht sng ven
bin nc c l loi Crocodylus porosus.
Ra bin thng sng vng nc nhit i, nhng i khi chng cng b dng
i dng cun ti vng b n i. Mt s loi ra n sa hoc c ngoi bin khi,
mt s loi khc nh ra xanh th n c bin vng nc cn, nhng tt c u di c
113

rt xa, vo tn t lin trng vo nhng khu c bit trn ct. T l cht khi
ang l trng rt cao do b cc sinh vt n tht t nhin n v c do con ngi. Nhng
ch ra con mi n cng b chim n trong qu trnh b ra bin v khi ra ti bin th b
c tn cng. Khi trng thnh th b con ngi bt n tht v ly mai lm trang
tr. Hin nay, ra bin c xem l loi gp nhiu nguy him nht v nhiu quc gia
thc hin cc bin php bo tn, trong c bin php ngn cm bt v nhp khu cc
ch phm t ra. Nhiu vng n Dng - Thi Bnh Dng v bin Ca-ri-b
c nhng n lc thu nht trng v gi chng n khi n, sau th xung bin,
ngi ta vn ang ch xem nhng bin php bo v ny c duy tr c dn s ca loi
ra khng.
Rn bin thc s l ng vt bin, hu ht u sng bin sut qu trnh sinh
trng. Ngi ta cho rng chng c lin quan n rn sng trong t lin lc a n
v tt c u sng trong vng nc m n Dng v Thi Bnh Dng. D
chng sng theo by ln, thc n ca chng l c nh nhng thc t ta vn cha bit
nhiu v c im sinh hc v sinh thi hc ca chng. Rn bin cc k c, bnh
thng khng hung d, nhng chng c th gy cht ngi. Chnh v iu ny, ngi
ta lo rng, knh o qua eo bin Panama c th lt rn bin vo vng nc m Ca-
ri-b v i Ty Dng. Hin nay chng b lp nc ngt trong knh chn li. Chng
l h hng ca rn h mang v thng sng vng n Dng v Thi Bnh Dng,
d dc bin pha ty ca Bc M cng c mt loi sinh sng. i Ty Dng v
a Trung Hi khng c rn bin. Rn bin c nhng chic rng nc c nh v rt
c, tuy nhin cng khng c nhiu ngi cht do b rn bin cn v c mt vi loi
rn li hin mt cch ng kinh ngc. Nhiu loi trng trn b, nhng mt s hon
ton sinh sng bin, rn con c sinh ra v ln ln lun trong nc.
Rn bin n c, nhng v cn ht th khng kh, nn chng phi ngoi ln mt
nc ly khng kh. Phi chng ln v di n tn gn ui. Mt s ln su n 150
m, nhng ni chung l 5 m. Mt vi loi c th di nc vi ting ng h v chu
ng rt tt iu kin thiu xy. Mt s ch ng by c trong cc hc san h v
sau chp ly con mi bng ming, trong khi mt s khc nm yn nh nhng ci
que v tn cng c khi c bi gn chng.
Ra bin, tng rt ph bin vng nhit i, nhng nay ang gp nguy him
do sn bt trc tip v mc vo li bt c, tm. Nm loi sng vng bin nc M
v Ca-ri-b (hnh 6-3) u l loi n tht, tr loi ra xanh n rong bin v c bin.
Kemp's ridley v ra Carretta n ng vt khng xng sng st y, loi ra lng
da n sa v i mi n bt bin. Ra terrapin c phn lng ng nh sng ca bin
c th b dc y v bt trai bng cch vn u xung rt nhanh. Cng c khi, n bi
rt kho bng cch p nhanh chn trc v pha sau, y mnh ln trc tn cng
cc con mi c linh ng cao hn nh cua. Ra xanh bi rt nh nhng vng nc
nng, tm n c bin. Ngc li, ra lng da n con mi l ng vt ph du c dng
jelatin.

114


Hnh 6-3: Ra bin c tm thy b bin nc M (Ngun: Hi ng nghin cu
quc gia, 1990).
6.1.5 ng vt c v (Mammals)
C ba b ng vt c v tin ho t t tin sng trn t lin v chng thch
nghi vi cuc sng di bin mt cch c lp. Ba b ny bao gm: (i). b c voi, c
heo v c porpoise (gn ging c heo); (ii). b hi cu, s t bin v hi m; v (iii).
b c nc, ln bin v b bin. Tt c u c chung c im ca loi c v, ngha l
c mu nng, con v u phi ht th khng kh. C voi v c Porpoise, thuc b
ng vt c v, bao gm nhm c rng Odontoceti v nhm c sng hm Mysticeti.
Tt c u c dng thun v c cu to rt ph hp vi cuc sng di bin. B c v
bao gm c voi v c porpoise.
B c v Cetacea bao gm khong 76 loi ng vt c v sng bin, nh c
voi, c heo, c heo m. T tin ca nhm ny l cc ng vt sng trn cn v di c
xung bin khong 55 triu nm trc. Loi ln nht l c voi c tm sng hm (hnh
6-4) v chng to nn mt phn b ring (Mysticed) vi khong 10 loi. Ging nh
loi c mp c kch thc ln nht, hu ht cc loi c voi v i ny ch yu l n
ng vt ph du, lc thc n qua mt tm sng hm chuyn bit. Tm sng hm trng
ging nh ci bn chi mc t vm ming c hai pha v thc n c tm sng hm
lc li, lu lu li c li ly i. C voi lng g v c voi lng xm cng c th bt
c nhng n c ln, nh c thu hay c trch. C voi xm ht thc n l ng vt
sng y.
Mt vi loi c voi c tm sng hm ln (nh c voi xm, c voi lng g) di c
rt xa theo ma, thng vo ma ng chng kim n vng nc nhit i v chuyn
v cc cc vo ma h. Cc loi nh hn khng di c xa nh th, nhng chng cng di
chuyn khi ngun thc n hay iu kin vt l thay i.
Phn b Odonticed bao gm 66 loi c v khc, tt c u c rng v c mt l
phun nc duy nht thay v l 2 nh c voi sng hm. Phn b ny bao gm cc loi c
115

voi cn li, c heo v c porpoise (hnh 6-4). C voi c rng l loi n tht rt kinh
khng bin, chng n mc v c, mt vi loi cn n c c voi khc hay hi cu v s
t bin. Khng ging nh c voi sng hm, chng khng n cc ng vt sng trn
mt v c th ln su xung n vi trm mt. C nh tng gi k lc v kh nng ln
su trong s cc ng vt c v bin; ngi ta cho rng n c th ln xung c
2200 m tm kim nhng con mc khng l. Mt s loi c rng sn bt mi bng
cch nh v bng m thanh, chng pht ra cc nhp m v nghe nhp phn m v t
nht c mt vi loi cng hp tc vi nhau dn mi v bt mi.
Mt s nh khoa hc cho rng loi c v ni chung v c th l ring loi c
nh tng, n mt lng ln hn c lng con ngi nh bt c trn ton th gii. V
d, t nm 1979 n nm 1982, 18 loi c v tiu th ht khong 46.000 ti 460.000
tn hng nm vng bin Georges Bank, ngoi khi pha ng bc ca M, trong khi
tng hi sn nh bt c cng vng ny l 112.000 - 250.000 tn. a Trung
Hi, ni mc l ngun thc n quan trng ca ngi, c tnh ng vt c v n mt
lng mc gp 2,3 ln ngi n. V trc thi im lng c voi b gim i 1/10 do
con ngi sn bt, c voi sng hm n n 190 triu tn tm hng nm, tc l ln hn
hai ln tng lng hi sn do con ngi nh bt trn ton th gii. T nhng con s
ny, ta c th thy rng cng khng ngc nhin khi nhng ng dn coi ng vt c v
nh l k cnh tranh vi h v c v mc.
T xa xa, ngi Eskimo sn cc loi ng vt c v bin, nhng sn bt
c voi bt u sm nht ngoi khi vng bin Bc u, t nm 800 n 1000 trc
cng nguyn v sn bt c voi tr thnh cng nghip mang li li nhun cao vo gia
nhng nm 1700 v 1800. C voi c khai thc ch yu ly du (thp n) v c
voi sng hm c dng lm cng o qun ca ph n; tht c voi khng c coi
trng lm (tr Nht).
C kh ho tng, tu b tc cao v cc tu ngm c s dng rng ri t
u nhng nm 1900 khin cho lng c voi gim ng k, v e do n nguy c
tuyt chng ca nhiu loi. Thm ch sau khi Hip hi c voi quc t (IWC) c
thnh lp nm 1946, lng c voi b sn vn tip tc tng ln, ti 65.000 con trong
nhng nm 1960. Lng khai thc trc y v c tnh s c cn li hin nay ca 13
loi nh bng 6-1. Trong s ny, 9 loi c a vo danh sch ang gp nguy him
t nm 1970. Vi kh nng sinh chm v thi gian trng thnh di loi c voi
ln, phi mt hng chc nm mi c th phc hi li c lng c mt i. Ch c loi
c voi chn l vn tng, n c kch thc tng i nh khng bao gi b khai thc
nhiu v lng c ny pha nam tng do lng c voi sng hm gim, nguyn nhn l
v hai loi ny cnh tranh vi nhau v tm. Nm 1986, IWC ng ra lnh cm sn
c voi nhm mc ch thng mi trong 5 nm, vi mong mun phc hi li s c
ang gp nguy him; lnh cm ny c ph chun vo nm 1991. Tuy nhin, cc bin
php ny khng bo v c cc loi c v nh, cng khng lm gim c s lng
c b cht do cc nguyn nhn khc. Li tri, li ti v cc dng c nh bt khc
bt hng chc nghn con c heo v porpoise hng nm; t nm 1971 n nm 1987,
116

hn 6 triu c porpoise cht do b mc vo li ti c ngi ta ging ra bt c ng
vng bin nhit i pha ng Thi Bnh Dng. Cc loi c v sng ven b hoc ch
bi vo chu cnh mi trng sng b nhim v hu hoi ang cng ngy cng
tng. Khng nghi ng v vic mi trng nhim ang e do c tm trng (mt loi
c voi) vng ca sng St. Lawrence pha ng Canada v e do c c heo nc
ngt nhiu vng.

C voi xanh
C voi lng xm
C voi "Right"
C voi "Sei"
C voi lng cong
C voi xm
C voi
sng hm
C voi
c rng
C voi hnh mi chai
C voi trng
C voi dn ng
C voi n tht
C nh tng

Hnh 6-4 So snh kch thc ca c voi sng hm v c voi c rng
B c v bin th hai gm hi cu, s t bin v hi m. Nhng loi tng t
nhau ny thuc loi chn mng (b chn mng), tc l 4 ngn chn ca chng c
dnh vo nhau bi mng da. Ngc vi c voi, nhng ng vt ny sng mt phn
trn t lin hoc trn nhng tng bng ni, chng t tp y n v ngh ngi.
Ngi ta tm thy 32 loi chn mng tt c cc bin trn th gii, c mt loi sng
h Baikal, nhng hu ht u sng vng nc lnh Nam cc v Bc cc. Thc n ch
yu ca chng l c v mc, nhng hi m cng s dng nanh o ng vt thn
mm v ng vt y khc sng y bin. Chng thng sng v di chuyn theo by
v mt s loi di c kh xa.
6.1.6 Chim bin (seabirds)
Chim bin l chim di chuyn mt khong cch rt xa trn bin v thng kim
n cc o xa ngoi khi hoc vng ven b. Chim bin c mt tt c cc i dng
v bao gm rt nhiu loi khc nhau. Chng c mt tuyn mui thi lng mui
117

n vo ra khi c th. Chim bin c th l chim cc khng bit bay cho ti chim fr-gat
c cuc sng hon ton trn khng, c th l loi ch n cc ng vt ph du nh ti
nhng loi n c ln, nh chim b nng. Mt s ch sinh sng mt vng nht nh,
trong khi c nhng loi di c ti hng nghn kilmt. Chim bin c tui th rt cao,
loi chim albatrosses c th sng n 50 nm. Hu ht chim bin tr mng bin, u
c s tin ho lm mt kh nng i vng vng trn mt t. Nhiu loi sng thnh n,
c khi ln ti hng ngn con, v c nhng n chim nhn bin gm hng triu i.
Chim bin thng sinh sn nhng ni cch xa mi nguy him v thng l nhng
o xa s hot ng ca con ngi. Chim bin gm 4 nhm chnh (bng 6-1 v hnh 6-
5).
Nhm chim cnh ct khng bay c. Cc ngn chn ca chng chnh l cnh
trc ca t tin bit bay tin ho thnh. Chng sng thnh by Nam cc v vng
nc gn Nam cc, vi s lng c th l vi i cho n hng ngn.
Nhm chim hi u P-tren v h hng ca chng nh albatrosses, p-tren,
shearwater v chim p-tren c kh nng ln v c cc l mi pha ngoi, rt hu ch
trong vic nh hi con mi. Loi chim albatrosses c cnh di n hn 3 mt v c
th bay ln rt gii, li dng c nhng lung gi n nh cc i dng pha bc.
Chng thng sng thnh by, n cc loi chim khc. Chim hi u rt c chuyn n c
con hay ng vt ph du, chng c cc dim ging nh lc mi bn ming chuyn
n ng vt ph du. Chng c th sng thnh n c vi nghn con, nhng c khi cng
ch vi con, mt s loi c thi quen di c rt xa.
Nhm B nng v h hng ca chng, thng c trng lng tng i ln v
gm nhiu loi c lng trang tr vi mu sc nh nhng. Chng thng sng vng
nhit i, nhng cng c loi sng Bc Cc v Nam cc. Mt s loi khc nh chim
chin (chim free-gat), li sng rt gn b.
Nhm Mng bin, nhn bin, v chim anca cho n nay l nhm chim bin a
dng nht. Chng c th sng thnh n hng triu con, nhng sinh sn th theo nhm
vi nghn con. Loi mng bin n c trch, vi nhiu loi c h hng gn gi vi nhau
sng khp ni trn bn cu Bc v trng cc vng dc b bin v o. Nhn bin
th nh hn, sng gn vi bin hn l mng bin v rt a dng vng nhit i, mc
d cng c nhng nhn bin sng ngay st vng Nam cc v Bc cc.
Ging nh loi b st v loi c v bin, chim bin c tin ho t cc loi
sng trn t lin. Hin nay c khong 260 - 285 loi, ph thuc vo cch nh ngha
v loi chim bin; chng chim khong 3% s loi chim tn ti trn tri t. Nhng
loi chim sng gn lin vi bin c chim anca, hi u p-tren, tt c u c vi i din
sng trong t lin, vng c nc ngt v sng bin n 50 - 90% cuc i ca
chng. Mt vi loi khc sng nh thc n bin nhng li khng bit bi.
Cc loi chim bin c nhiu cch kim n khc nhau v thc n cng rt a
dng. iu ny c th hin s khc nhau ca m v cnh cc loi. Mt s loi
nh nhn bin m dp, mng bin, hi u th bay l l ngay trn mt nc, s khc nh
118

b nng, n bin th lao su xung hn di nc bt ng vt ph du, mc, hay c.
Chim cnh ct, chim cc, hi u rt c, ch ng tm mi di nc, dng cnh hoc
chn bi. Chim cnh ct c th ln su xung n 250 m, nhng ni chung hu ht
cc loi chim bin u kim n lp nc trn b mt. Tc ng do chim bin kim n
trn mt n i sng sinh vt bin thng b b qua, nhng c th tc ng l rt
ng k.

Cnh ct vua
Hi u
Chim chin
"frigate"
Mng bin
lng en

Hnh 6-5: Mt s loi chim bin: (a) cnh ct vua, (b) hi u, (c) chim chin
"frigate" v (d) mng bin lng en.
Mc d chim bin c mt khp ni, nhng thng nhng n ln sng gn
nhng ni c iu kin thun li cho sinh sng v thc n di do. Nam cc c ti
hng triu con chim cnh ct (su loi), thc n ch yu ca chng l tm hoc c v
mc. Tng t c nhng n chim vi s lng rt ng sng cc o vng bin
ngoi khi pha Ty Nam M. Ngoi bin, chim thng tp trung sng dc vng b
bin, ni c cc iu kin thun li cho pht trin sinh hc. Cn vng nhit i c
iu kin sinh trng km hn th chim bin t hn nhiu. S phn b ca chim bin c
th phn nh nhng thay i theo ma mi trng bin v mt s loi c tp qun di
c rt xa ti ni c sn ngun thc n hay ni c iu kin thi tit thun li cho
sinh sn.
6.1.7 C (fish)
C l chi ph bin trong qun th sinh vt bin, c khong 20.000 loi c sng
cc va san h, ca sng, hm ni di bin, nhng vng nc nm khong gia v
y bin su. Rt kh phn bit loi c sng y v c sng trong tng nc gia v
tnh di ng cao ca chng v v c rt nhiu mi trng sng di bin. C l nhm
ng vt c xng sng bin phong ph nht v ln nht. Chng c phn thnh ba
lp nh sau:
Lp Agnatha, bao gm cc loi c nguyn thu nht, c khng c xng hm,
c mt v c mt myxin. Nhm ny tin ho t nhng loi c mt t 550 triu
nm trc Cambrian, nhng hin nay ch cn khong 50 loi.
119

Lp Chondrichthyes, c c mp v h hng nh c ui, nhng loi ny u
thuc loi c nhm; c im ni bt ca chng l c xng sn v khng c vy. y
cng l nhm c c, xut hin ln u tin vo khong 450 triu nm trc v hin nay
cn khong 300 loi.
Lp Osteichthyes, bao gm c c xng thc s. y l nhm ph bin nht v
rt a dng, vi khong hn 20.000 loi sng bin. Chng c tin ho t 300 triu
nm trc.
1)- Agnatha: Thn ca c mt v c mt myxin thon di v trng ging nh
ln, khng c vy. Bao quanh ming l mt b phn dng mt v hu ht u sng
nh n tht cc loi c khc. C mt myxin c vo xc ng vt cht n phn
bn trong; cn c mt l loi k sinh, chng bm vo c khc bng phn ming v
cn vo trong n phn mm v ht cht dch ca c th con mi. Tt c cc loi c
mt myxin u sng bin, cn mt s loi c mt sng vng nc ngt.
Thm ch mt vi loi c mt d sng bin nhng khi cn nh chng sng trong
sng, n cc loi ng vt khng xng sng nh v c khi c c. Sau khi ln ln,
chng chuyn ra sng ngoi bin.

Hnh 6-6: Mt s loi c c xng vi hnh dng khc nhau
2)- Chondrichthyes: C mp thng c xem nh l mt loi c bi nhanh v n tht
hung d nht bin, chng n c nhng ng vt rt ln, nhng thc ra, cn c nhiu
loi khc cng hung d nh vy. Nhng c mt nghch l l, cc chi ln nht ca
nhm ny li l nhng ng vt rt hin lnh ch n cc loi ph du, l c mp
basking v c mp voi vi chiu di thn tng ng vi tng loi l 14 m v 16 m. C
hai loi u c rng nh v lc ng vt ph du bng nhng ci mang c bit c
tin ho. C ui v c ui skate c phn thn dt v hu ht sng vng st y.
120

Hu ht u n cc ng vt y (c bit l cc loi gip xc, nhuyn th v ng vt
da gai), nhng mt s cng n c v c ui manta loi ln th n cc loi ph du.
C mp v c ui thng th tinh bn trong v kh nng sinh sn khng cao,
ch rt t trng. Hu ht cc loi c mp v tt c cc loi c ui u con. C
ui trng nhng khoang c bo v tt, thng gn vo mt cht nn, c con
sau s n ra trong vng vi tun hoc vi thng.
Nhu cu v tht v vy c mp ngy cng tng, chu , vy c mp c coi l
mn n c sn. Hng nghn con c mp b mc vo li tri dng nh bt mc.
S c mp ang gim rt nhanh nhiu ni, chng hn nh vng bin pha ng ca
Florida, ni ngh nh bt hi sn rt pht trin.
3)- Osteichthyes: Nhiu loi c c xng sng nhng mi trng bin khc nhau v
do vy m chng to nn mt nhm khng ng nht v cu trc, sinh hc v c im
sinh thi.
Nhng loi c c xng quen thuc nht l loi c ngi ta nh bt vi mc
ch kinh t, do m c im sinh hc ca nhng loi c bit ny cng c bit
n nhiu hn. Cc loi ny n nhiu loi thc n khc nhau, tu thuc vo kch thc
ca chng, ni sinh sng v ngun thc n cc thi im khc nhau. Mt s ch n
sinh vt ph du, mt s khc n c, hoc c khi n c hai. Nhng loi c ng o ny
ng bc thp trong thp dinh dng; chng gm c trch, c mi cm hay c sacin
v c trng. Tt c u n ch yu l ng vt ph du, ngoi ra c trng trng
thnh cn c th n trc tip to ct kt thnh mng ln. Nhng loi c ln, nh c
tuyt, c meluc, c p-lc, khi mi n n nhng ng vt ph du c nh. Khi ln
hn mt cht na, chng chuyn sang n cc loi ph du ln hn (chng hn nh tm
vy) v sau khi hon ton trng thnh th n c. Nhng loi c c xng ln
nht sng ngoi khi u n c, chng hn nh c ng, c nhng. Mt vi loi nh
c tuyt, c phin, c meluc, kim n c tng nc gia, y bin v c th bt c
hay ng vt khng xng sng di y. Loi c demersal thc s hon ton sng
gn y hoc ngay trn y bin, trong khi mt s loi khc (nh c bn) ch n sinh
vt y (trai, giun, v cc loi gip xc), mt s khc li n nhng con c nh hn.
S phong ph ca ngun thc n cho c bin l khc nhau ty thuc cc yu t
vt l. Trc s bin i ngun thc n theo ma c th bit trc c mt s loi di
c ti nhng vng nht nh khi c nhiu mi. Chng hn, c ng Thi Bnh
Dng di c ti vng bin ngoi khi ni c nhiu n cua sng theo ma. Tuy vy,
nhiu loi c, lng thc n bin i c th dn ti s thay i v s sng v pht trin
ca chng, iu ny c phn nh qua thay i lng c nh bt c gia cc nm.
6.2. Cc phng php thu thp mu
Loi v kch thc ca li r vng nc trung gian c thit k sao cho c
th bt c sinh vt tri ni t su ny ti hn 1000 m (Harrisson, 1967). Li
Isaacs-Kidd l mt chic ti hnh nn di, ming c ng knh 8 m v c mt b phn
121

gi cho li nm bn di mt nc khi ko. Ngi ta thng lp thm mt my
ghi su v su trong lc ko c iu khin qua mt my nghe trong nc bng
cch gn mt b phn pht ra m thanh rt nhy vi p sut, tc l tn s nhp m
thanh s thay i theo su. Cng c loi li c ci tin c hai u, m c
iu khin bng van p sut.
Li r vng nc trung gian (Baker v cng s, 1973; Clarke, 1969) c
pht trin s dng v nh bt su hn 1500 m, c mt ci ming m v ng
bng b phn iu khin t xa (xem hnh 6.7). Khi ming ng, li c h xung
su cn thit, c xc nh bng tn s nhp rung ca ca b phn pht m thanh rt
nhy vi p sut gn trn li. Sau , ming li m ra bng mt c cu iu khin
bi mt tn hiu m thanh t trn tu. Lc gn cui, mt tn hiu m thanh th hai t
tu s lm ng ming li trc khi ko ln. Mt loi ci tin khc t loi ny gm
hai li c iu khin bng m thanh trong cng mt khung chung, li pha trn
nh c kch thc mt li 0.32 mm nm trn li c kch thc ln hn 4.5 mm v
ming m 8 m
2
. Cng c mt b phn iu khin nm trn khung o su, dng
chy v nhit ri truyn cc d liu ny v tu bng tn hiu m thanh.

B phn nhn m thanh v o su
Thanh c nh ming li
Hp s
Thanh c nh ming li
Thanh trt gn
vo y li
Thanh nh
v y bin
Li
B phn thu mu

Hnh 6-7: Li r cc sinh vt vng nc trung gian, c RRS Discovery s dng.
Li c m v ng bng cc tn hiu m thanh truyn i t tu. su ca li
c xc nh bi tn s nhp rung ca m thanh pht i t mt b phn nh v
su. (a) Li c ng li h xung (b) li m (c) li c ng ko ln.
6.3 cu hi v tho lun

122

123
Chng 7
Sinh vt y

7.1 PHNG PHP PHN LOI
Vng bin ngoi khi cha ng nhng mi trng sng rt a dng, khc nhau
v su, nhit , nh sng, mc ngp (triu hay vng chu triu mt phn) v kiu
tng nn khc nhau. Nhng tng nn cng l ni c ng ca cc loi sinh vt khng
chn nh ng vt chn t v trai c - nhng loi nh c mt v tr nht nh trong
sut thi k trng thnh v nhng nn ny cng c nhng khe nt, vt ln l ni n
np an ton trnh loi n tht ca cc ng vt c kh nng di chuyn cao. Cc tng nn
mm (nh bn, st, ct) va cung cp thc n va l vt bo v cho cc ng vt o.
Do mi trng sng y a dng hn m s loi ng vt y (khong 160.000)
nhiu hn rt nhiu ln s ng vt ph du (khong 3.700).
Vng y nh nht l vng trn mc nc triu, mt vng nm bn trn mc
nc cao nht do triu v ch b ngp khi c bo. vng b bin dc, vng ny b nc
bn ln t nhng con sng v v th i khi c gi l vng bn to (splash zone).
B bin thoi l vng c c bin, mt vi loi thch nghi vi cuc sng ni tip gip
gia bin v t lin.
Vng nm trong bin triu (littoral hay intertidal zone) l vng c bin
trn v di gii hn bi mc nc triu, do vy khi triu cao th ngp, khi triu thp th
l ra. Phm vi ca vng ny ph thuc vo a hnh v bin triu. To y v thc
vt ph du l thc n ca ng vt n c y bin v nhng sinh vt ny l ngun dinh
dng cho cc loi n tht.
Vng di mc nc triu (sublittoral hay subtidal) nm t mc nc triu
thp ti ra ngoi ca lc a, su khong 200 m. Mt phn ca vng ny thuc
vng st mt (euphotic zone), nhng s loi thc vt y gim dn ti khng khi tin
ti nhng vng su hn. Tng nn him dn v thay vo l tng nn mm. Vng
di mc nc triu chim khong 8% tng bin chu ngp.
Nhng mi trng sng vng y cn li nm bn di vng euphotic. Vng
bin su tnh t su 200 m ti 2000 hay 3000 m (gii hn thp hn cha c xc
nh), v chim khong 16% tng bin ngp. Vng abyssal (su thm) t 2000 hay
3000 m ti 6000 m, cho ti nay, chnh l vng sinh hc ln nht, chim khong 75%
mi trng sng y. Vng ny c nhit thp hn 4
o
C. Vng su nht ca bin l
t 6000 m xung ti khong 11000 m t c bit n nht do kh tip cn, v c t loi
sinh vt sng c nghin cu.

7.1.1 Thc vt y (Benthic plants)

124
C rt nhiu loi thc vt bin sng y hoc sng trong trm tch vng
bin cn. Tt c u thuc vng euphotic; ngha l chng u thuc vng trong bin
triu v vng di triu cn.
Cc qun x sinh vt bin ch yu vng nm trong bin triu l cc loi ht
kn (loi cy c hoa), sinh trng nhng vng c trm tch, cho php r c th mc
c. Cc qun x ny gm c m ly ngp mn vi nhiu loi cy c kh nng chu
mn cao; nhng vng s vt ca sng v c bin vng ngp triu. Tt c loi thc vt
y loi ln ny c kh nng sinh trng cao, nhng chng cng cha mt t l kh ln
cc thnh phn khng th tiu ho c i vi hu ht cc ng vt. Do vy m chng
to nn mt lng tng i cc cht phn hu v c dng triu a n nhng vng
bin khc, gp phn vo ngun dinh dng cho nhng vng bin ny.
Thc vt bin c kch thc ln bao gm c to bin ln- loi sng rt mnh
nhng b bin c vng n i. Mt vi loi to xanh (nh Halimeda) v to
(Lithothamnion) c kh nng ht cac-bo-nat can-xi vo trong m, do vy m chng
khng b cc loi n c tiu th. To san h cng ny mc nh mt lp cn vi bm
bn ngoi , v c hay san h v chng gp phn to nn lp trm tch vi.
To mt (nh Ectocarpus) sinh trng trn b mt ca cc loi thc vt ln hn
(c bin, to). Chng thng c dng si, v c cc loi n thc vt bin rt a
thch.
Sinh vt sn xut y him nht l to n bo sng trn cc ht ct, hoc to
nn mt lp trn b mt lp bn. Cc loi thc vt c kch thc nh ny bao gm to
ct, to xanh l cy xanh da tri v to roi. Cc sinh vt ny thng rt pht trin v
d c kch thc b, chng vn l ngun dinh dng quan trng cho nng sut c bn
nhng vng nc cn.
Cc loi thc vt y khc nhau sng vng chu mc triu khc nhau v s
phn i ny mt phn c quyt nh bi kh nng hp th bc sng nh sng ca
chng. mt bin, ngi ta thu c quang ph hon chnh ca nh sng nhn thy
c, nhng nhng bc sng khc nhanh chng c hp th v phn tn trong ct
nc.



Hnh 7-1: Mt vi v d v to bin (a) Enteromorpha (ti 500 mm); (b) Ulva (ti
250 mm); (c) Alaria (ti 2 m); (d) Chondrus (ti 150 mm); (e) Gigarfina (ti 200 mm);
(f) Delessefia (ti 250 mm); (g) Fucus vesiculosus (ti 1 m). (a) v (b) l to xanh; (c)
v (g) to nu ; v (e) v (f) l to .

To lc thng ch sinh trng vng nc cn v sc t ca chng hp th c
sng ngn v sng di. To nu v to cng c cha cht dip lc, nhng chng c
nhng sc t c bit khc to nn mu sc ring bit. So vi to lc, to nu (to b,
to thch y) sng kho hn vng nc su hn, sc t chnh ca chng, fucoxanthin,
hp th nh sng c bc sng ngn tt hn. Hu ht to (v d Gigartina) sng
vng di mc nc triu; sc t ca chng ng vai tr quan trng trong hp th
nh sng xanh lam xanh lc xuyn su trong nc. Tuy vy, cng c nhng ngoi l.
7.1.2 ng vt y (Benthic Animals)
ng vt y c chia lm hai loi da vo ni sng ca chng, lin quan ti
tng nn: Infauna v Epifauna. Trong Infauna l loi c ton b hoc mt phn
cuc sng ca chng trn tng nn, bao gm trai, giun v cc loi khng xng sng
khc (hnh 7-2). Chng thng sng tng nn mm v rt a dng, phong ph vng
di mc nc triu. C t sinh vt loi ny tng nn cng. Loi Epifauna (hnh 7-3)
to nn mt nhm ng vt ln hn sng y bin, khong 80% ng vt y thuc
loi ny. Mt vi loi tiu biu nh san h, sao bin, bt bin, trai, chn t. Epifauna c
mt cc loi tng nn nhng c bit pht trin tng nn cng, sinh trng mnh
vng nm trong bin triu v cc rn san h. C th thm vo mt loi th ba
na, l ng vt cng sng gn vi y bin nhng thnh thong bi ngay bn trn
125

nn bin; nhng ng vt ny (nh cua, c dt nh c bn), to nn mt loi gi l
epibenthos.

Hnh 7-2: Mt s infauna (a) Hydrobia, mt loi c; (b) hang ca Pygospio, a
polychaete; (c) hang ca Corophium, mt loi gip xc chn hai loi; (d) Arenicola a
polychaete; v trai (e) Cardium, (f) Macoma, (g) Scrobicularia, and (h) Mya.

Hnh 7-3: Mt vi loi sinh vt y
126

127
Cng c cch phn loi ng vt y theo kch thc. Trong trng hp ny,
kch thc ca ng vt c lin quan n kch thc ca sng dng lc ng vt ra
khi trm tch. Cc loi sau y bao gm ht tt c mi kch thc ca ng vt y:
a)- H ng vt ln - Macrofauna (hay macrobenthos): nhng ng vt ny ln
hn 1 mm. y l nhng ng vt y ln nht, bao gm sao bin, trai, san h,
v.v.
b)- H ng vt c trung bnh - Meiofauna (hay meiobenthos): Nhng ng vt
ny c kch thc t 0.1 n 1 mm, thng sng trong ct hay bn. Loi ny
gm ng vt nhuyn th, nhng con giun rt nh, vi nhm gip xc nh (bao
gm chn cho y) v mt s ng vt khng xng sng t quen thuc.
c)- H ng vt c nh - Microfauna (hay microbenthos): Kch thc ca loi
ny nh hn 0.1 mm. Loi ny gm cc ng vt nguyn sinh, loi c t (Hnh
7-4).
a)- H thng phn loi v sinh hc ca chng
Cc qun x y bao gm mt tp hp rt a dng cc ng vt y. Nhiu loi
sinh vt bin khng h c h hng sng trn cn hay nc ngt v l nhng loi ng
vt him. Bng 7.1 cho thy mt s qun x ng vt y ph bin v c minh ho
cc hnh 7-5, 7-7.
Ngnh ng vt nguyn sinh y c bit n nhiu nht l foraminifera. C
hng ngn loi, v chng l mt phn c bn trong cc ng vt y c trung bnh v
nh, c bit trong trm tch bin su. Mc d l cc sinh vt n bo, nhng khng
nht thit phi l c kch thc nh, mt s di n 25 mm. C c epifauna v infauna,
v ni chung chng n cc loi to ct y vng nc nng, cc loi nguyn sinh
khc, cht thi ra v vi khun tt c cc vng su khc nhau. Mt trong nhng
loi mi c pht hin thuc loi ny l Xenophyophoria (hnh 7-5a), pht trin rt
mnh nhng vng b bin dc. y l loi sinh vt n bo ln nht c ng knh
ti 25 cm, nhng ch dy c 1 mm.
ng vt a bo c bn nht l bt bin (hnh 7-5b), mt s vng chng pht
trin rt mnh. Ngi ta cho rng chng tn ti t cui thi Precambrian (hn 600 triu
nm trc), v nhm c xa ny hin nay c ti khong 10.000 loi, hu ht sng
bin. Chng c t tn theo bn cht sng vng dc (ngnh vng dc), v nhng
hang hc ca bt bin chnh l ni c ng cho hng ngn ng vt b nh nh giun v
gip xc. Tt c cc loi bt bin u khng c b phn bm, ngha l chng gn kt
vi nhau v khng di chuyn c. Chng lc thc n bng cch sinh ra cc dng cun
cc ht vt cht vo. L ca bt bin ng vai tr nh ci sng, ch cho php cc ht
nh nht i qua v gi li cc ht bng cc t bo dng roi c bit.



Hnh 7-4 Mt s v d v ng vt c nh
(ng vt nguyn sinh c lng mao vi nhiu hnh dn khc nhau)
Thc n bao gm phn ln cc loi vi khun, sinh vt ph du v cc ht b
phn ra nh. Phn khung xng ca bt bin c cu thnh t cac-bo-nat can-
xi hay cc gai silic. V c gai cng v kh nhn, nn bt bin c rt t nguy him
loi tr mt vi loi c sng cc rn san h, mt s loi c v thn trn. Chng
sinh sn v tnh v hu tnh v c kh nng phn tch t mt nguyn th ban u.

Hnh 7-5: Mt s ng vt y c bn
128

129
(a) Xenophyophore n bo; (b) Bt bin; (G) C chn ngng bin; (d) giun ng; (e) Giun
nhiu t; (f) Giun dt; (g) Enteropneust hemichordate; (h) eGhiurid; (i) sipunculid; v (j)
ng vt mang ru. (Tt c cc loi trn c kch thc tnh bng mm)
Ngnh sa lng chm c rt nhiu loi sng y. Nhm ny cng c lch s tin
ho lu i v hin nay c nhiu loi sng hu ht cc mi trng bin. Hu ht loi
sng y ngay trn tng nn, nhng cng c mt vi loi sng trong lp ct bn.
Trong ngnh Lng chm, lp thu tc bao gm cc loi thu tc tp on (hnh
7-3), c to nn bi hng lot cc c th c cu trc v chc nng khc nhau. Lp ln
hn l San h, vi hn 6000 loi, bao gm cc loi c chn ngng bin v cc loi san
h cng vi mt vi dng t quen thuc hn. Chn ngng bin (hnh 7-5c) l qun th
vng nm trong bin triu v di mc nc triu ph bin nht, nhng n cng
c tm thy vng bin su hn 10.000 mt; l cc ng vt th cng, ng
knh t 1 cm ti hn 1 m.
Giun y thuc mt s ngnh khc. Cc loi giun trn ging nh si ch (ngnh
giun trn) to nn mt nhm ng vt bin (v c trn cn) phong ph nht v ph bin
nht, mc d hu ht cc loi ny sng n mnh trong lp trm tch mm. Ngnh
Phylum Nemertea (hnh 7-5d) gm khong 600 loi giun di, tt c u c mt ci vi
di c th ln ra ngoi bt mi. Chng sinh trng mnh cc vng bin n i hn
l bin nhit i, v thng vng cn. Giun dt sng t do (ngnh Platyhehninthes)
(hnh 7-5f) sng trong bn, ct, bn di v v trai c hoc sng trn rong bin,
nhng t khi chng sng theo tp on ln. Ngnh Sipuncula (hnh 7-5i) cn gi l
giun c lc, l loi giun khng c t c chiu di t 2 mm ti hn na mt. Nhiu loi
lm trong ct hay bn, s dng ci vi ln ca mnh to nn ng i xuyn qua
lp trm tch; mt vi loi khc sng trong cc hc hay hc san h, thm ch sng
trong cc v trai c rng. Thc n ch yu ca chng l cc cht phn hu. Ngnh
Echiura (hnh 7-5h) ni chung t nhiu tng t vi Sipunculids v kch thc v tp
qun sinh sng, nhng khc ch l chng c vi ln khng c kh nng co gin, dng
tm kim thc n trong bn ct.
Ngnh Pogonophora (hnh 7-5f) cng pht trin mnh vng nc su hn,
xung ti 10.000 m. Nhng loi giun khng chn bm ny vn nhng ng dai v di
ra bm vo tng nn cng.
C hn 10.000 loi thuc ngnh Annelida, lp Polychaeta, to nn mt nhm
a dng v ln nht trong s cc nhm giun bin. Polychaetes (hnh 7-5e) l cc loi
giun phn t vi nhiu chn, gi l chi bn. Chng c chiu di t vi mm ti 3 m. V
mt sinh thi hc, polychaete c th c phn thnh loi bi mt cch tch cc trong
lp nc st y bin hay lm t trong ct, bn v loi sng trong ng hay hang vnh
vin. Nhm ny, bao gm c epifauna v infauna, gp mt phn ln trong sinh khi
y nhiu mi trng sng.
Ngnh Hemichordata gm enteropneusts (hnh 7-5g), hay gi l giun u,
thng sng vng nm trong gii hn mc nc triu v vng thu nhit bin su

130
v cc rnh. Loi ln nht c bit n c chiu di ti 1.5 m, nhng hu ht u c
kch thc nh hn nhiu. Nhiu loi sng trong hang hc trong bn ct, mt s khc
di chuyn mt cch chm chp trn b mt lp trm tch hay to thnh nhng lp bo
nho trn nn cng.
Cc chi ca ngnh nhuyn th gm hn 50.000 loi sng bin, trong s c
c l mt loi quen thuc v cc loi thn trn hay c bin khc (lp chn bng) v cc
loi trai hai mnh v. Mc a dng cao trong ngnh ny th hin s c mt ca
loi nhuyn th tt c cc tng su ca bin, v c trn v trong lp bn ct, c
i din tt c cc bc dinh dng v trong mi qun x sinh vt y.
ng vt da gai (ngnh da gai - hnh 7-6) l nhng loi ch c bin. C
khong 5.600 loi, c phn thnh 5 lp:
- Lp Asteroidea bao gm khong 2.000 loi sao bin sng t vng trong bin
triu xung ti su khong 7.000 m.
- Lp Ophiuroidea bao gm khong 2000 loi c sao hoc sao bin (hnh 7-6c).
Nhng bc nh chp t di bin su cho thy cc loi ny ph ln y bin, chng
kim n lp trm tch, thc n l nhng ng vt sng hay cht, hoc nhng cht
hu c l lng.
- Lp Echinoidea bao gm khong 800 loi nhm bin gai (hnh 7-6d) v loi dt.
- Lp Holothuroidea (500 loi) bao gm cc loi da chut bin di, c tn gi ny
do chng ging nh qu da chut. Cc loi da bin ny n c thc n l lng v
thc n lng ng, chng sng trong ct cn n c bn.
- Lp Crinoidea l nhm da gai c xa nht, hin nay cn khong 650 loi, ch yu
l hu ty bin v sao lng. Sao lng (hnh 7-6a) thng sng su 1.500 m,
mc d sng bm vo nn bin, chng vn l nhng loi c kh nng di ng cao
bng cch b hay bi.
ng vt ru (hnh 7-7b), thuc ngnh Ectoprocta. Ging nh thu tc, ru l
nhng ng vt sng theo tp on v khng c chn bm, to nn lp ph mng trn
nm trong ranh gii mc nc triu, ph ln v trai c hay c nhng b mt vt nhn
to. Mt s loi sng su hn 8.000 m.
ng vt tay cun (hnh 7-7c) to nn ngnh tay cun gm t hn 300 loi
ng vt bin rt ging vi loi nhuyn th ch c v can xi hai mnh (ng knh 5-
80 mm), tuy nhin phn thn c bn ca chng th hon ton khc. Hu ht sng
su nh hn 200 m gn lin vi tng nn cng. Tuy vy, mt s loi ng vt tay cun
sng cc t theo chiu thng ng trong ct v bn v mt s loi sng su
5.500 m.
ng vt c v (hnh 7-7d) l ng vt y c h hng vi larvacean v salps.
Chng c hnh tr trn khng chn bm v thuc ngnh Urochordata, lp Hi tiu.
Hu ht cc loi c v thng thng u cng, nhng cng c nhiu loi sng theo tp

on v sinh sn v tnh. Hi tiu thng c tm thy vng nm trong bin mc
nc triu, bm vo , v c, cu tu hay cc cht nn cng khc, nhng chng cng
sng su hn 8.000 m.
Gip xc c t xut hin nhiu y bin. Cc loi c kch thc trung bnh
bao gm ostracods, cyclopoid, harpacticoid copepods. Chn kim l mt nhm sinh
trng rt mnh, chng trn hoc o xuyn qua trm tch mm. Cng thuc lp c
kch thc trung bnh ny l Tanaids (hnh 7-7e). Cc loi gip xc nh ny (chiu di
< 2 mm) c thn hnh tr mnh hn, v thng o l hoc sng trong ng, c mt
mi su, su nht ti hn 8.000 m. C khong 350 loi, nhng ti nay c im sinh
hc vn cha c nghin cu r.

Hnh 7-6: Mt s v d v ng vt da gai y
(a) Sao lng (Mt loi hoa hu bin); (b) Da chut bin (Mt loi da bin); (c)
brittle star (Mt loi sao bin d v dng ui rn); v (d) nhm bin (Tt c cc loi
trn c kch thc tnh bng mm)

131
Gip xc y c ln thng gm c ng vt chn ging isopods (hnh 7-7f) v
ng vt chn hai loi amphipods. Chn ging thng c mnh dt, di 5 - 15 mm, tuy
nhin cc loi sng su hn thng di ti 40 cm. Ngi ta thng quan st thy chn

ging chy rt nhanh trn trong vng bin triu, nhng mt s loi th o hang
c mt nhm lm ng hm trong g. Phn ln trong s khong 4.000 loi ny l loi
n tp v xc thi. Chng sng vng bi cn v trong cc mng rnh.
ng vt chn t l gip xc khng chn bm duy nht. C khong 800 loi,
bao gm mt s ln k sinh trn ng vt bin khng xng sng. Chng ging nh
tm, sng trong lp ph canxi pha ngoi. Hu ht sng vng nc cn, nhng cng
c vi loi sng su hn 7.000 m. Loi chn t sng t do kim n bng cch qut
nhp nhng phn chn trong nc. Chng thng bm vo y tu, b mt bn tu.
Gip xc y 10 chn bao gm cc loi cua, tm thng thng, v nhm ny c
i din sng c trn mt v trong lp trm tch y bin. C rt nhiu loi sng vng
nc cn, nhng cng c loi sng su 5.000 -6.000 m. Nhm ny bao gm c
loi n tht, loi n tp v n xc thi. Nhiu loi trong nhm ny l ngun thc n rt
quan trng ca con ngi, v cng vi cc loi nhuyn th, chng to nn mt ngnh
cng nghip nh bt hi sn c v.

Hnh 7-7: Mt s v d v ng vt y
(a) scaphopod; (b) Tp on ru; (c) ng vt tay cun c cung; (d) ng vt c v;
(e) tanaid; (f) ng vt chn ging; v (g) Con sam. (Tt c cc loi trn c kch thc
tnh bng mm)
C t ngnh ng vt bin khc khng c cp y, v chng c t loi
hoc do chng thng khng sng y. Mt vi trng hp, chng ta vn thiu s
hiu bit v vai tr sinh thi hc ca mt s nhm ng vt no ch v thit b thu
thp mu hin nay khng bt c chng. Nhiu loi mi (cng nh nhiu h mi)
132

c pht hin trong vng 25 nm qua, ch yu l di bin su, v hy vng danh
sch ny vn ang tip tc c ko di.
7.2 PHNG PHP THU THP MU
C mt s ngnh khoa hc nghin cu v y bin; chng hn nh ngnh i
dng hc, a cht hc, c sinh vt hc v sinh hc bin. Mi mt ngnh u s dng
nhng thit b ring c thit k thu thp thng tin cn thit cho ngnh ca mnh,
nhng c rt nhiu s trng hp gia cc lnh vc v khoa hc bin, m d liu ca
ngnh ny c th cng rt hu ch cho ngnh kia, do c nhiu dng c thit b
nghin cu y bin c th cung cp thng tin cho ngnh sinh hc bin, y ch
cp n mt s thit b thng dng nht. Nhng thit b di y gm c hai loi: (i)-
thu thp mu trm tch, v (ii)- thu thp sinh vt y. S khc bit gia hai loi cng
khng phi l r rng lm, v mu trm tch cn cha c cc sinh vt nh v thit b
dng bt cc sinh vt y cng c th s mang theo c trm tch.
Hnh 7-8: Thit b ly trm tch theo thi gian.
Trm tch i vo phiu hnh nn v ri xung ng thu mu c cha fomandehit. Sau
mt vi ngy, khay ly mu s t xoay v ng thu mu tip theo s nm ng vo v tr
ca phiu hnh nn.
bin su, lng cht hu c c nh hng n thnh phn ca qun x ng vt.
Khi lng v bn cht ca cc ht sinh vt tch t xung trong nm c th o c
bng thit b ly trm tch. Thit b ny c a xung cc su khc nhau v
trong sut c nm hay thm ch lu hn. Cc thit b hin i lm vic t ng, trong
c nhng ca vo xoay trn (hnh 7-8). Mi ca m trong mt s ngy, thng l 7
133

134
k thut chp nh nh k c
s dng rng ri trong nghin cu Dng i d
i
khi c
hn nh y bin nhn
ung trn ton cu, nng sut ca
thc vt y chim cha ti 10% tng nng sut c bn bin.
hng chic ti c
mt l
g dn theo qu trnh sinh trng v
s ti s
c xc nh v vic ly mu c thc hin lin tc o s thay i sinh
khi, th
ngy, sau ng li (Lampitt, 1996). K thut ny v
ng ton cu (Ducklow v Harris, 1993).
a)- Cc php o nng sut c bn ca thc vt y
Nng sut ca thc vt y c kch thc ln thng c c tnh qua kt qu
thu hoch v tng khi lng ca cy, o theo n v cac-bon trn mt n v din tch.
nhng vng n i ni cy ci pht trin theo ma kh r rng, thng thu hoch khi
cy giai on c sinh khi cao nht; i khi thu hoch c lng sinh khi ca l
hay cung mc ln, nhng cht hoc bin mt trc v sau khi thc hin o c.
Khi c mt s thay i nh theo ma trong sinh khi, s tng trng ca l hay cy
th bit c bng cch c cc l ln cy; nhng ci l ny s cao dn ln cng
vi s pht trin ca cy v ngi ta o c s tng trng ny qua chiu cao .
Thc vt sng gn lin vi vng y c li th l c ngp trong dng nc
chy ri mang theo cht dinh dng ho tan v lin tc i mi. Nng cc cht dinh
dng tng ln vng nc ven b v t l cht dinh dng c ht vo thng cao
thc vt y. Thng thng s sinh sn trn mt n v din tch ca to vng ny ln
hn so vi thc vt ph du. Mc d s sinh sn quang hp di y pht trin mnh
hn so vi thc vt ph du nhiu vng ven bin, ch c mt p
nh sng cho cc loi thc vt sng y. Ni ch
b)- Ly mu v php o sn xut ca ng vt y
C th thu thp c ng vt y bng cc thit b ly mu khc nhau. vng
nc c su nh hn 30 m, quan st trc tip, m v thu thp bng bnh kh p ca
th ln l k thut hiu qu cho php th ln thu bt k mt loi mu tng nn no v
cho php nh gi kh tt iu kin t nhin, bao gm c s phn b khng ng u
ca sinh vt y. su nm ngoi gii hn an ton ca th ln, gu xc thng
c s dng thu mu cc qun x sng trong trm tch mm. Nhng thit b ny
ly c mt mu y theo ng lng cn thit t c th thu c ng vt
sng bng cc loi ry. Li vt l nhng thng cha c trang b n
i th kh chc chn v c ko trn y bin; chng c thit k thu bt
ng vt y, nhng chng khng cung cp thng tin v khi lng.
Lng ng vt y thng xuyn c o bng s ng vt trn mt n v
din tch (m
-2
). Sinh khi y c biu din theo gm
-2
, mt cch l tng th nhng s
liu ny th hin theo s gam carbon hay s gam trng lng kh, nhng cng c th l
theo s gam trng lng t ( hon chuyn). Sinh khi thay i lin tc v php o
nng sut th cp c tnh n iu ny. Sinh khi tn
n xut ca cc c th trong mt qun th hay qun x; sinh khi hin c b mt
i do b n hay nguyn nhn gy cht khc no .
C th phn loi theo thi k sinh trng. Nu sinh khi tng thi k sinh
trng
ngi ta c th tnh c nng sut trong khong thi gian gia cc ln ly
mu.

135
g
Agassiz
nh 7-9). N c u im l rt d s dng v chm y bng mt no cng c.
Ming ca li c gi trng thi m bng mt khung kim loi v gn vi li c
mt dy thu bt cc sinh vt nh. Ngi ta ch vic ko li st di y.

Nu s sinh sn khng gii hn trong mt thi gian c nh th khng th o
mt nhm ng vt n l. Khi c nhiu lp thi gian chng nhau, vic nh gi trc
tip sn lng th cp ca sinh vt y tr nn kh khn hn. Trong nhng trng hp
nh vy, nu c th th nghin cu trong phng th nghim xc nh t l tng trn
ca mi loi theo kch c bt k no . Qun th ny sau c chia ra thnh nhiu
loi kch thc m khng tnh n thi k sinh trng v sn lng c tnh bng
cch nhn s ng vt c trong mi loi kch thc vi t l sinh trng ca loi .
Mt loi li thng c s dng trong lnh vc o c sinh hc l
(h

i Agassiz

7.3 cu hi v tho lun

Hnh 7-9 : L

Chng 8
DNG NNG LNG V
CHU TRNH KHONG CHT

8.1 mng li thc n
8.1.1 Mng thc n vi khun
a)- Khi nim v chui thc n (Food Chain)
Chui thc n l mt qu trnh tuyn tnh biu th trong cc sinh vt tiu th
cc sinh vt khc trong cng mt mi trng. Cc chui thc n cho bit s chuyn i
nng lng v cht hu c thng qua cc bc dinh dng khc nhau ca cc sinh vt.
Mi bc dinh dng c to nn bi cc sinh vt cng nhn nng lng theo
mt cch tng t nhau. Chui thc n bin bt u t cc thc vt ni; cc sn phm
s cp t dng ny to nn cc cht hu c t cc cht v c v to nn bc dinh
dng th nht. Cc loi ng vt ni n thc vt (c) s dng trc tip to bin to nn
bc dinh dng th hai v chng c xem nh sinh vt tiu th ban u. Cc bc dinh
dng tip theo c to bi cc loi ng vt ni n tht cc loi ng vt n c, v
cc sinh vt tiu th bc cao hn th n cc sinh vt tiu th bc thp hn. Tng s cc
bc dinh dng s thay i theo tng vng v s loi trong qun x sinh vt. Bc dinh
dng cao nht l cc ng vt trng thnh khng b n tht bi ng loi ca chng;
cc sinh vt bc cao nht gm cc loi nh c mp, c, mc v cc loi ng vt ln
khc. Tng sinh khi c to ra trong tt c cc bc dinh dng cao hn trn mt n
v din tch v trn mt n v thi gian c gi l sn lng th cp. Cc nghin cu
ng hc dinh dng nhm kim tra cc yu t nh hng ti s chuyn ho nng
lng v vt cht gia cc bc dinh dng v cc yu t iu chnh sn phm th cp.
Cc nguyn t nh ni t, cc bon, pht pho c kt hp vi nhau trong cc
thnh phn hu c ca cc m ng thc vt, chng c mt chu trnh khp kn thng
qua chui thc n (hnh 8.1). Vi khun tiu th cc cht thi v cc m ca cc sinh vt
cht. S phn hu ny to ra cc dng v c ca cc nguyn t cn thit cho cc sinh
vt t dng. Dng nng lng c hng duy nht. Mt phn nng lng b tn tht ti
mi ln chuyn ho ti bc dinh dng tip theo v rt nhiu ho nng b tiu th trong
dng hu c c chuyn sang nhit nng v b tiu hao do h hp, khi cc bon hu c
chuyn thnh CO
2
. Nh mt chui lin tip, tng nng lng s gim ti mi bc dinh
dng v iu ny to ra gii hn v s bc dinh dng trong mi qun th.
136


Hnh 8-1 : Chu trnh khong cht v dng nng lng trong h sinh thi bin.
b)- Hiu sut sinh thi (Ecological efficiency)
S chuyn i t bc dinh dng ny sang bc tip theo l khng hon ton, tc
l khng phi tt c sn lng ca mi bc dinh dng c chuyn ho hon ton
sang bc tip theo. c tnh sn lng ti bc cao nht ca chui thc n (v d sn
lng c to ra) ta cn phi bit nng lng mt i. Chng c th c tnh t hai yu
t sau:
1. Khng b n. Mt s i tng ca bc dinh dng trnh b tiu th bng
cch trn thot, khng chp nhn, hoc khng c gi tr. V d thc vt ni c
gai ln hoc c c cc ng vt ni trnh khng n, hoc thc vt ni qu
nh hay qu ln khng ph hp lm mi cho cc sinh vt khc.
2. Khng c kh nng chuyn ho. Mt s i tng ca thc n khng c kh
nng chuyn ho cho c th pht trin.
Chng ta c th thit lp cn bng thc n nh sau:
I = E + R + G (8-1)
trong I l lng b n, E l lng bi tit, R lng tiu th do h hp, v G l lng
s dng cho tng trng; G c th c chia thnh pht trin c th v ti sn xut. Mt
cn bng nh vy thng c lp theo dng n v nng lng hoc calo.
S pht trin c th thng chim mt phn nh trong cn bng trn. Khng
phi tt c thc n u c th c tiu ho v ng ho, m mt phn b bi tit. Mt
phn nng lng nhn c trong thc n b tn tht khi h hp, v th n khng c
chuyn sang bc dinh dng tip theo.
137

S chuyn ho khng y trong chui thc n c tnh bng hiu sut chui
thc n. E c nh ngha l phn nng lng c c t mt bc dinh dng chia cho
phn nng lng cung cp cho bc dinh dng . Hiu sut tng trng l t l thc
n ch yu c dng cho s tng trng c th thay i trong khong 30 45%,
nhng hiu sut chui thc n thng thp hn nhiu, thng ch khong 10%. Cc h
thng ph du v cao c hiu sut chui thc n cao hn.
Hiu sut nng lng chuyn ho gia cc bc c gi l hiu sut sinh thi
(E) v n c nh ngha l phn nng lng c c t mt bc dinh dng cho
chia cho nng lng cung cp cho bc dinh dng . Hiu sut sinh thi rt kh o,
n ch c xc nh gn ng t hiu sut chuyn ho (E
T
) nh sau:

1
=
t
t
P
P
T
E (8.2)
trong Pt, l sn lng hng nm trong bc dinh dng t, P
t-1
sn lng hng
nm trong bc dinh dng t-1. Trong phng trnh ny, sn lng c th c xc nh
c dng nng lng hoc sinh khi (v d gram cc bon). i vi s chuyn ho nng
lng gia thc vt ph du v ng vt ph du, E
T
s bng sn lng ng vt n c
chia cho sn lng s cp. Ti bc tip theo, hiu sut chuyn ho s l sn lng
hng nm ca cc ng vt tiu th bc hai (ng vt ph du n tht) chia cho sn lng
hng nm ca sinh vt n c.
i vi mt vng cho, s bc dinh dng c th gp i cng vi s tnh
ton nh lng nng sut s cp v t d tnh nng sut th cp (P) trong bt k bc
dinh dng no da theo phng trnh sau:
P = B.E
n
(8-3)
Trong B l sn lng s cp hng nm, E l hiu sut sinh thi, n l s
chuyn ho dinh dng (bng s bc dinh dng - 1). Kh khn c bn trong vic p
dng phng trnh ny l chnh xc ca cc gi tr trong cng thc hiu sut sinh
thi v s cc bc dinh dng.
Li thc n l s phc tp hn ca cc tng tc thc n. Hnh 8.2 cho thy
li thc n ca vng bin Bc, cui cng l c trch. Cc quan h thc n kh phc tp,
nhng chng c th c n gin theo chui thc n tuyn tnh nh pha tri ca
hnh v, n xem xt cc loi chnh thng qua cc chu trnh hu c. Mi loi biu th
mt bc dinh dng c nh ngha nh mt loi hoc nhm loi m tt c nui dng
mt loi hoc cc loi khc. Cc loi trong mt bc dinh dng cng c th b tiu th
bi cc loi bc dinh dng cao hn.

138


Hnh 8-2: Li thc n c cu trc t cc quan h nui dng ca c trch bin Bc,
trong sut cc thi k sng khc nhau. Chui thc n n gin ca c trch c v
bn tri (theo Russell-Hunter, 1970.)
c)- Li thc n vi sinh vt (Microbial food web)
S ti to cc cht dinh dng trong bin l mt trong nhng tng tc quan
trng gia cc bc dinh dng cao hn v thp hn. iu ny c thc hin bi vi
khun v cc sinh vt nguyn sinh, n tng gp i vi chui thc n truyn thng c
hnh thnh bi thc vt ph du - ng vt ph du - c (hnh 8.3). Cc dng ht vt cht
trong bin c hnh thnh do xc cht ca cc ng thc vt ph du v sinh vt tri
ni, hoc qua sn phm cht lng ng khc nh lt xc ca cc loi gip xc, rng
lng ca cc loi ng vt v phn hu bi vi khun. Cc loi vi khun cng c th s
dng cc cht hu c c hnh thnh bi cc qu trnh bi tit ca ng vt v o thi
ca thc vt ph du nh hnh 8.4, trong trng hp ny vi khun chuyn ho mt cch
hiu qu cc cht dinh dng b phn hu sang sinh khi.
S lng vi khun trong vng nc mt ca i dng thng khong 5
triu/ml. i khi chng t ti 100 triu/ml nhng ni giu dinh dng v t cc loi
tiu th vi khun. Ti cc vng nc su, s lng vi khun c th nh hn 1000/ml.
Ni chung s lng vi khun trong bin c iu chnh do vic sn mi ca cc sinh
vt ph du, c bit l cc sinh vt n bo khc nhau, mt s ng vt ph du loi ln
cng tiu th vi khun. Trong s cc sinh vt ph du, cc loi ng vt hnh roi l tiu
th vi khun nhiu nht. Cc loi d dng thng c mt khong 1000/ml. Tuy
nhin khi s lng vi khun bt u tng ln th cc sinh vt cng tiu th vi khun
tng ln nhanh chng. Cc ng vt ph du nh l nhng cu ni quan trng trong vic
chuyn ho sn lng vi khun sang mc dinh dng cao hn, ng thi chng to ra
139

ngun thc n cho cc sinh vt ln hn, c th l cho cc loi gip xc. Ni chung, mt
s loi nh loi chn kim khng th n trc tip vi khun v b phn lc ca chng
qu ln gi li cc vt nh b.

Hnh 8-3 S m t s nhn i chui thc n ca c (T ng vt ni n c n
tht) v vng kn vi sinh vt (vi khun v sinh vt n bo). ng nt t ch s gii
phng cht hu c b phn hy (DOC). Cc ng vt nguyn sinh tiu th vi khun,
sau n lt chng b thiu th bi ng vt ph du ln hn.

Hnh 8-4 : Nhng con ng chnh chuyn ho v ti to cc cht hu c trong bin.
140

Thng thng qu trnh xoay vng nh m t trn c biu th hnh 8.4, n
cho thy hot ng vi khun trong bin lin kt cht ch vi cc li thc n. Nhng
thay i trong ngun thc vt ph du thng ph thuc theo thi gian v khong cch
do s thay i sinh khi vi khun. Trong qu trnh tng trng theo hm s m ca
thc vt ph du, vi khun c th sng khng cn chuyn ha cht hu c b phn hu,
chng c to ra nh mt phn ca qu trnh chuyn ho ca qu trnh tng trng
ca thc vt ph du. Cui thi k pht trin mnh nht ca thc vt ph du, khi m to
bt u giai on gi, c mt s tch lu cc vt cht t ngun gc thc vt ph du v
s gii phng cc cht chuyn ho c tng ln. C th l ti thi im vi khun c
th s dng cc ngun nng lng ny tng ln v to ra s bng pht mnh m. T
thc vt ph du cng pht trin mnh. Mi quan h gia thc vt ph du v vi khun
nh hng ti phn b thng ng ca vi khun ph du. S lng cc i ca vi khun
thng xut hin ni thoi ho t bo, ni m vt cht thc vt tch lu do lng ng
t vng nc mt. , s phn hu do vi khun gp phn hnh thnh cc lp c t
xy nht trong cc vng nc tnh. Ni chung, ngi ta c tnh rng bng vic s
dng cc vt cht thc vt hoc cc cht tit dch hu c ho tan cho s pht trin m vi
khun c th dng ti 50% hoc hn lng cc bon c c nh bi quang hp.

Hnh 8-5: S thay i trong cc mc dinh dng ca cc h sinh thi ph du,
phm vi t cc mi trng i dng ti cc mi trng ven bin v ti cc mi trng
dinh dng cao xo trn mnh (Theo Landry, Helgolinder wissenschaffen
Meeresuntersuchungen, v. 30, pp. 8-17, 1977.)

141

8.2 cc chu trnh khong cht (C, N, P)
Tt c cc nguyn t hnh thnh nn cht hu c u c ti sinh, nhng vi
khong thi gian khc nhau. Qu trnh chuyn ho cc cht hu c tr li dng nguyn
t thng c xem nh s khong ho. N din ra trong ton b ct nc t mt cho
ti y bin, ni c nhiu vt cht b ri t cc lp nc pha trn tch lu li. S phc
hi cc khong vt c th din ra tng i nhanh (trong mt ma) vng mt nc,
hoc chm hn nhiu trong trng hp cc vt cht kh b ri lng xung v tch ng
y bin.
Cac bon dioxit (CO
2
) v cc cht dinh dng c tr li cho vic ti s dng
nh thc vt ph du. Chu trnh l khc nhau trong cc khu vc ni khng c xy ho
tan t do. iu kin thiu xy c lp bn ct b mt y bin v mt s vng bin
c bit nh bin en su khong 200 m n y bin bi v s trao i nc v
pha trn vi bin a Trung Hi gn k b gii hn bi a hnh y bin. Di cc
iu kin ny, s suy gim vi khun din ra do cc vi khun k kh s dng xy t cc
gc sulphate v nitrate. Loi o xy ho ny to ra cc hp cht rt yu nh methane,
hydrogen sulphide, v ammonia. V cc hp cht ny nhiu ho nng, cc nhm khc
ca vi khun c th s dng nng lng ny to ra cc bon dioxit v to ra cht hu
c mi. Qu trnh c nh cacbon t CO
2
to ra cc hp cht hu c bng vic s dng
nng lng nhn c t xy ho cc cht v c (v d nh nitrite, ammonia, methane,
sulphur) c gi l ho tng hp.
Trn quan im sinh thi, vn quan trng nht ca s ti sinh cc cht trong
bin l tc ti cc cht dinh dng hn ch sinh trng c ti to. Trong cc
cht dinh dng c cung cp tc th trong bin, nitrate (NO
3
-
), phosphate (PO
4
3-
), v
cc cht ho tan nh (Si(OH)
4
) thng thy nhiu nht trong vng tp trung pha di
mc bn bo ho cn thit cho pht trin cao nht ca thc vt ph du. S hn ch silic
dioxit nh hng u tin ti cc sinh vt s dng nguyn t ny to xng, chng
gm c to ct v cc loi to khc trong s cc ng vt v thc vt ph du. Chu trnh
silic tng i n gin v n ch gm cc dng v c; cc sinh vt s dng silic ho
tan to xng ca chng v xng s phn hy sau khi cc sinh vt cht.
8.2.1 Chu trnh Cac bon (Carbon cycle)
Cac bon khng ging nh nit, pht pho hay nhiu nguyn t khc, cc bon
khng bao gi c mt trong bin vi s lng c hn. Tuy vy, chu trnh cc bon (hnh
8.6) c mt s tnh cht c bit, bao gm c cc qu trnh vt l v sinh hc.
Cac bon dioxit vo trong i dng t kh quyn v n ho tan nhiu trong nc.
Nu nng CO
2
trong nc bin ph thuc hon ton vo p sut phn t ca CO
2

trong kh quyn (0.3 ml/l), vo nng tng i ca CO
2
trong nc v khng kh,
vo nhit v mn ca nc, th khi lng CO
2
trong nc bin s rt thp.
Cc ion ny c to thnh cc dng cac bon dioxit v chng biu th cho s
bn vng nht ca cac bon dioxit trong nc. Trung bnh c khong 45 ml CO
2
trong 1
lit nc bin. Bi cc phn ng cn bng ho hc nu trn, s tng vt cc ion
bicarbonate v carbonate hot ng nh mt b cha CO
2
, khi CO
2
t do c loi b
142

bi quang hp th phn ng chuyn sang tri v s tng vt cc dng ion gii phng
nhiu hn CO
2
t do; v vy thm ch khi c nhiu quang hp, CO
2
khng bao gi l
yu t hn ch i vi sn lng thc vt. Ngc li, khi CO
2
c gii phng bi h
hp ca thc vt, vi khun v ng vt, th cc ion bicarbonate v carbonate c to
ra nhiu hn.
Ch rng cc ion hydro c gii phng trong cc phn ng ho hc chung.
iu ny c ngha l pH ca nc bin c iu tit ln bi nng bicarbonate
v carbonate, v thng l 8 + - 0,5. Khi CO
2
c b sung vo nc bin nh qu
trnh khong ho v h hp th khi lng ion hydro tng ln v pH gim xung
(dung dch tr nn axit hn). Nu CO
2
b loi khi nc do quang hp th iu ngc
li xy ra v pH li tng ln. Nh vy nc bin hot ng nh mt dung dch m.
Mt s sinh vt bin kt hp cc ion can xi vi cc bon trong qu trnh vi ho
to xng. Canci cacbonat (CaCO
3
) c th c dng calcite hoc aragonite
(khong), sau dng d ho tan hn. Sau khi cht, xng chm xung v c phn
hy, trong trng hp CO
2
li c gii phng vo nc, v cng c chn vi
trong bn ct y, trng hp ny CO
2
c loi b khi chu trnh cc bon.
Chu trnh cc bon n gin c m t trong hnh 8.6 . Ni chung, CO
2
c
chuyn ho t cht v c sang cc bon hu c nh qu trnh quang hp ca thc vt
ph du. Sau c hp th bi cc mc dinh dng cao hn, v mt phn CO
2
c
ti quay vng nh bicacbonat v c trong khi mt phn b tn tht t b mt i dng
trong dng kh. Carbon dioxide c hp th ti b mt i dng v c to ra trong
ct nc bi qu trnh h hp v khong ho. Mt iu chc chn l cacbon dioxit
c hp th bin nhiu hn trong kh quyn.

Hnh 8-6 S c bn ca chu trnh Cac bon

143

8.2.2 Chu trnh Nit (Nitrogen cycle)
Chu trnh Nit trong bin (Hnh 8.7) kh phc tp v n xut hin nhiu dng
m khng d chuyn ho t dng ny sang dng khc. Chng bao gm nit phn t
ho tan (N
2
) v cc dng ion ca ammoniac (NH
4
+
), nitrit (NO
2
-
) v nitrat (NO
3
-
), cng
nh cc hp cht hu c khc nh ure (CO(NH
2
)
2
). Dng ch yu ca nit trong bin
l ion nitrate v dng ny thng c cc thc vt ph du s dng, mc d nhiu loi
cng c th s dng nitrit hoc ammoniac. Mt s t loi thc vt ph du c th ly cc
phn t nh nit hu c nh l axit amino v ure. Vn tc ti nit trong trng thi
n nh c to sn cho thc vt ph du c th gii hn nng sut s cp trong nc t
dinh dng quanh nm v trong nc m trong ma h.
S phc hi nit trong ct nc l do cc hot ng ca vi khun v bi tit ca
cc ng vt bin, c bit s bi tit amonia do ng vt ph du. Nh trong hnh 8.7,
s xy ho amonia thnh nit v sau thnh nitrat c coi l s nit rt ho; cc vi
khun iu chnh thay i ny trong trng thi ho hc c gi l vi khun nit rt
ho. Qu trnh ngc li ca vic to thnh cc hp cht nit b gim t nitrat xut hin
hu ht trong bn ct t xy v c gi l s kh nit; cc thay i ny c tin
hnh bi cc vi khun nit ho. Chu trnh nit cng gm c s c nh nit, trong
kh nit b phn hy c chuyn ho thnh cc hp cht nit hu c; qu trnh ny c
th c thc hin ch bi mt s t thc vt ph du. C hai loi nit hu c phn hy
(DON) v nit hu c phn t (PON) u l dinh dng cho s pht trin ca vi khun.
Vi khun ph hy cc protein thnh a xit amonia v amonia, v sau b xy ho
trong qu trnh nit rat ho. Cui cng s gii phng nit v c ho tan (DIN) lm cho
cc dng ny li c sn cho cc thc vt ph du tiu th. Cc dng khc nhau ca vi
khun trong chu trnh ny c th phc v bn thn chng nh ngun trc tip ca thc
n cho mt s ng vt ph du.

Hnh 8-7: Chu trnh Nit trong vng nh sng ca bin. S phc tho s quay
vng ca nit din ra trong vng nh sng v vng pha di ng dinh dng.
144

Hy ch n cc quan h ni ti gia nit v c ho tan (DIN), nit hu c dng
phn t (PON) v nit hu c ho tan (DON). ng dinh dng l ng ti c
s thay i nhanh chng nng cht dinh dng theo su.

Mt vn quan trng ca chu trnh nit trong bin lin quan n ngun nit s
dng trong sn phm u tin. Mt s phn ca sn phm s cp nhn c t nit ti
hi phc t vt cht hu c trong vng mt nc; mt phn khc nhn c t nit mi
t cc ngun bn ngoi vng nh sng (hnh 8.7 v 8.8). Nit mi l nit rt s
cp vo vng nh sng t vng di nh s xo trn theo chiu thng ng, nhng
n cng gm nhng lng nit nh hn vo do s c nh nit v t dng chy sng
ngi v ma. Nit c hi phc dng ban u amoniac v urea.
S so snh nit mi v ti to (xem hnh 8.8) l quan trng bi v ch ngun vo
lin tc ca nit mi c th xc nh tng kh nng ca i dng i vi sn lng
nh bt c bn vng (nh rng loi b c t i dng cng l loi b nit). Cng ch
nit mi th mi c th gip loi b s vt qu CO
2
c a vo i dng t cc
hot ng ca con ngi; trong trng hp ny, sn phm tng ln ca thc vt ph du
loi b nhiu hn CO
2
trong bin.

Hnh 8-8: So snh sn phm to thnh bi nit ti hi phc v nit mi. Kt qu
sn phm ti to ch t nit ti hi phc c cung cp trong vng nh sng do bi
tit ca sinh vt. Sn phm mi ph thuc vo nit c cung cp t bn ngoi vng
nh sng, trong ngun ch yu l nittrt chuyn ln t tng di. Trong trng
thi n nh, nit chuyn ln cn bng vi lng chuyn xung trong cc ht bn ct.

Trong hnh 8.9, t s quang hp (PQ l t s ca s mol xy to bi thc vt
ph du chia cho s mol xy b s dng) c dng d on s khc bit trong cc
qu trnh to sn phm bao gm hai loi nit. Ch rng sn phm c ti to da
trn nit phc hi c PQ (khong 1,2) thp hn sn phm mi (khong 1.8) lng ny
i vo bin t kh quyn v sng ngi, c bit da vo nit rat lp nc pha trn.
Nguyn nhn ca iu ny c tm tt hnh 8.9, cc gi tr khc nhau ca PQ
145

nhn c ph thuc vo loi nit ang c dng v con ng ho hc ang c
thc hin trong qu trnh quang hp. Nu ch vt liu cac bon hydrat ang c hnh
thnh, th s c mt lng xy c sinh ra cho mi mole cac bon dioxit (CO
2
) b s
dng. Tuy vy, cht lipid cng c hnh thnh trong qu trnh quang hp v v cc
lipid b gim nhiu hn cac bon hydrat, nn xy s b loi tr thm v PQ s khong
1,2 thay cho 1,0. Nu cc lng ln nit rat (ngun nit mi) b ly i v b gim
xung trong qu trnh hnh thnh protein, thm ch nhiu xy hn c gii phng
lin quan n CO
2
b s dng, v nh vy PQ s tng ln khong 1,8. Nu amoni (dng
nit ti sinh) c dng nh mt ngun cho sn xut protein bi cc thc vt ph du,
th khi xy l cn trong qu trnh v PQ s l 1,0 hoc nh hn. V vy s phn
chia nhanh chng qun th thc vt ph du, s dng nit rat, s c cc gi tr PQ tng
i cao. Cc gi tr PQ thp hn cho bit trng thi gim st ca nit, ging nh amoni
c ti sinh, ang c s dng trong qu trnh quang hp.

Hnh 8-9 Mi quan h gia cc cch quang hp vi t s PQ. Phng trnh c bn ca
quang hp nm gia; trong trng hp ny, ch c carbohydrate c to ra v PQ
= 1 (11 mol 0, c to ra t 1 mol CO
2
). Ch rng gi tr ca PQ thay i theo kiu
ngun Nit c s dng bi thc vt ph du t protein
8.2.3 Chu trnh Pht pho (Phosphorus cycle)
Chu trnh pht pho kh n gin trong quan h ho hc; khi pH mang tnh
kim trong bin, pht pht hu c tng i d thy phn tr li thnh pht pht v c
(l loi c sn cho thc vt ph du). Bi v pht pho chuyn ho nhanh chng thng qua
chui thc n, n him khi b gii hn trong mi trng bin. So snh vi silic v pht
pho, chu trnh ti to ca nit l mt qu trnh phc tp hn. Chu trnh pht pho tng
qut c trnh by trong hnh 8.10.

146


Hnh 8-10a Chu trnh Phot pho tng qut (Theo Kupchella v Hyfand, 1993).

Hnh 8-10b Chu trnh Phot pho trong nc bin

147

8.2.4 Ph dng (Eutrophication)
Ph dng c nh ngha l s gia tng qu mc lng dinh dng v cht
hu c, gy ra nhng hu qu cho mi trng. Cc cht dinh dng v cht hu c l
nhng yu t c bn ca cc h sinh thi. Rt kh phn bit gia tnh thay i t
nhin v cc nh hng nhn to. Trong kha cnh ny ph dng khc vi cc vn
gia tng t ngun vo do con ngi gy c hi. Cc h sinh thi khc bit mt cch t
nhin trong s tp trung dinh dng, v mt h sinh thi ny c th c trng thi dinh
dng cao hn h sinh thi khc. Ph dng thng c coi l vn cht lng
nc rt nghim trng, trong con ngi ng vai tr c bn v a vo cc h
sinh thi qu nhiu dinh dng, c th l nit v pht pho. Ni chung, c th thy rng
cc ngun nit tng hp vo bin l yu t quan trng nht gy ra ph dng
(Smetacek v cng s, 1991) nhng iu ny c th khc nhau tu theo khu vc.
Cc ngun dinh dng:
Ngun thi khng c x l hoc x l c th chy trc tip vo bin. Cc
con sng cng l cc ngun im quan trng ca dinh dng v cc hp cht hu c,
chng gin tip vo cc vng ven bin. S dng t, quy m dn s v cc thit b
x l nc thi trong lu vc s quyt nh cht lng nc cc con sng chy vo
bin, v chng li nh hng ti cht lng nc cc vng ven bin. Tuy vy, cc con
sng khng phi l cc ng ng b ng v s chuyn ti ca sng c kt hp vi
cc qu trnh sinh hc v ho hc. V d, s tng trng ca thc vt ph du trong qu
trnh chuyn ti ca sng s thay i s phn b dinh dng v phn hu cc cht. Cc
nng cao ca nit v pht pho trong cc con sng kch thch thc vt ph du pht
trin. Trong ma xun iu ny c th dn ti s bng n ca to ct, ti nit v
pht pho l phong ph, c th lm suy gim cc cht khng tan ca silicat. V vy,
bin s nhn mt ngun vo vn giu nit v pht pho nhng ngho silicat. iu ny
dn n kt qu l suy gim sinh khi to ct v tng to roi. Hn na, hnh thc trong
nit i vo cc vng ven bin b nh hng bi mc x l nc thi v cc qu
trnh xy ra trong bn thn dng sng. Cc du hiu cho thy iu ny c th c hu
qu i vi thnh phn ca cc qun x to pht trin trong cc ngun nc t sng ra.
Ngoi cc ngun im ny, cn c cc ngun dinh dng quan trng t cc
ngun khuch tn. V d dng chy t t nng nghip hoc cc hu qu ca xi mn,
v lng ng t kh quyn.
nh hng ca cc ngun dinh dng:
S tng qut hnh 8.11 tm tt chui cc s kin lin quan n ph dng,
m t mt s nh hng ca ph dng trong cc ngun nc ven bin v cc iu
kin kh hu. Mt yu t rt quan trng l thi gian hi phc ca nc, n ph thuc
vo hnh thi ca nc nhn v s phn tng ca dng nc.

148


Hnh 8-11. Cc mi quan h trc tip v gin tip gia ph dng v s gia tng hm
lng cht dinh dng u vo , dn ti, (+) = gia tng; (-) = suy gim

So vi nc ngt trong cc h, s xo trn ca nc bin kh ln to nn s
cung cp xy ln v nh hng tt ti cc i dng v nhiu vng ven bin. Tuy vy
nhiu mi trng sng ca bin (cc vnh, cc bin kn) c c trng bi tc ti
sinh chm ca nc. V vy, cc vn ph dng ni bt nht trong cc vng bin
bn kn, v d nh bin Adriatic, nh hng ca ph dng do kch thch s pht trin
to v tiu dit vi khun ca cht hu c. Hn na, nc m c th cha t xy hn
nc lnh lm gia tng cc vn lin quan n suy gim xy trong nc.
Nhn chung, nh hng u tin ca s gia tng dinh dng l s gia tng sinh
khi v nng sut ca thc vt ph du. Thc t s gia tng nh dinh dng thng
c nh gi tt v n c th lm tng sn phm cc loi thy sn c gi tr thng
mi. Tuy vy, s gia tng nng sut v sinh khi thc vt ph du s dn ti gim lng
nh sng vo to y v tng bin ng v hm lng xy. Hm lng xy cao s
xut hin trong thi gian ban ngy, khi nh sng cho quang hp, nhng vo ban
m th sinh khi ln ca to s tiu th mt khi lng ln xy cho h hp v phn
r. S dao ng ln ca mc xy v suy gim xy l mt kt qu tt yu.
149

Ngoi gia tng sinh khi ca to, cn thy s suy gim tnh a dng cc loi v
tng xut hin cc bng n v to trong nc. Cc s bng n ny c th gy tr ngi
cho li thc n, chng c th sn sinh ra mt khi lng ln dng bt v lng ng
trn cc bi bin hoc thm ch c th gy c hi v dn ti cht hng lot cc loi
thy sn.
8.3 cu hi v tho lun

150


150

Chng 9
CC QUN X SINH VT Y
Cc qun x sinh vt y thuc loi mi trng bin phong ph nht, c bit
nhng mi trng sng ven b trong vng triu. Chng nhn c s cung cp dinh
dng giu c, chu nh hng c cc ngun dinh dng t t lin v cc ngun t
thc vt ph du ca bin c. S giu c ca cc mi trng sng ny lm cho chng tr
thnh ni nui dng v l vn m ca nhiu loi sinh vt di c, c bit l c, gip
xc, v chim. S cung cp dinh dng giu c v s thm ving ca cc loi n tht ti
vng ny thng dn ti cc chu trnh gia tng dn s ca cc loi mi, sau dn s
b ph v do cc loi n tht.
9.1 Mi trng vng triu
9.1.1 Thy triu v vng triu (Tides and intertidal zone)
Thy triu ln xung theo chu k ca mc nc bin vi mt khong thi gian
cho trc v chng c sinh ra do tng tc ln nhau gia lc hp dn ca mt trng,
mt tri vi tri t, v lc ly tm do s quay ca tri t v mt trng. Ti hu ht cc
b bin, thy triu ln xung hai ln mt ngy (bn nht triu) to nn cc vng ngp
triu c hai ln ngp nc mi ngy. Mc d vy, do cc iu kin vt l c th, mt s
vng ch c nht triu nh vnh M xi c.
Bin thy triu ln nht (triu cng) l vo ma xun, chng xut hin 2 ln
trong thng khi m tri t, mt tri v mt trng nm trn mt ng thng. Bin
thy triu nh nht xut hin khi cc hnh tinh ny khng nm cng ng thng.
nh triu l mc nc cao nht trong ngy v chn triu l mc nc thp nht. Phn
m rng ca vng ven b trong bt k a phng c th no cn c vo dc ca
ng b v bin triu, mt phn chng c xc nh bng hnh thi ng b.
Bin triu thay i t mc c th nhn bit r cc vng nh bin Ban Tch n
rt ln (ti 15 m nh vnh Fundy pha ng Canada).
Vng thy triu l vin st b ca y bin gia mc triu cao nht v thp
nht. Phn trn ca vng triu d xc nh trong cc vng nc tnh, nhng ti cc
vng b bin sng ln ln nhiu so vi gii hn bnh thng ca thy triu th li kh
xc nh, v iu ny m rng min khng gian sng ca nhiu loi vng triu.
Mt iu quan trng, l tnh cht ca vng triu phn vng theo chiu thng
ng, s xut hin ca cc loi chim u th trong cc di ngang ring bit. V d
(hnh 9-1), cu trc cc vng b bin n i. T cao nht n thp nht cc vng
l (a) vng a y en, (b) vng cy da cn, (c) vng cn hu chim u th, v (d) vng
cc loi chim u th khc nhau tu thuc vo tng a phng.


Hnh 9-1: nh hng ca sng ln sinh vt vng thu triu British Isles
EHWS: Mc nc cao nht khi triu cng (extreme high water spring tide); MHWN:
Mc nc trung bnh khi triu km (meanhigh water neap tide). Trong vng nc tnh
lp gia ca vng triu (bn phi ca biu ) loi sinh vt sinh sng ch yu l to
bin. Nhng ti vng chu nhiu tc ng ca sng hn, loi sinh vt ch yu li l
hu v trai. Ti lp trn ca vng triu rn n c l sinh vt chim u th. Vng c tr
chu tc ng rt mnh ca sng (phn bn tri biu ) c m rng theo chiu su.

Tm quan trng ca vng triu: Nh chng ta bit s quan trng ca lp hn
hp nc ven b, vng triu (tr im ni gia hai th gii sinh hc l t lin v
bin) to nn gii hn trn ca chng. Ti giao ct ny s ri lon ca nc c cc nh
hng mnh lit nht, tuy nhin cc chuyn ng ny cng to nn mt s giu dinh
dng rt ln, chng d dng c phn b bi dng ri. Khi nc nng, s trao i
vi kh quyn v s thm nhp ca nh sng vo nc kh thun li. Cc h sinh thi
c bit chim lnh cc mi trng ny gm:

Cc di san h: nng sut t ti 1000- 2000 g C/m
2
/nm.
Cc h sinh thi to: nng sut c th t ti 1500 g C/m
2
/nm hoc hn 3 kg
cht kh/m
2
/nm.
Cc h sinh thi b : chng rt a dng, cc ng vt thn mm 2 mnh v
sng bm vo trong khi cc loi chn bng sng khe hoc hang ng; cc
loi gip xc, c, giun v nhiu nhm ng vt khc cng c tm thy y.
Cc loi trai c kh nng bm sng cao nht v c th chu c sng. Trai
thng tp trung vi s lng ln ti cc b n su hn 10 m. Sinh khi
ca chng trong t nhin t ti 100 kg/m
2
dc vng b hoc vng su hn.

151


152
Cc h sinh thi bi bin: Nn bi bin ko di ti mt su vi mt thng ra
bin v c c trng bi nng sut sinh hc thp v thiu n nh v t nhin.
Cc ca sng: Do chng chim vng giao thoa gia nc ngt v nc mn,
cc khi nc ny thng tn ti ring r vi nhng nm mn xm nhp bn
di nc ngt. Thc t l cc ca sng hnh thnh cc h sinh thi a dng v
phc tp, v d c sinh vt ph du c bit l cc loi chu c khong thay i
rng ca mn. S khc nhau v mn v s c mt ca nc ngt thng
mang nhiu ngun dinh dng v cc loi t t lin, chng lm cho cc ca
sng giu c v nng sut cao, nhng cng rt c khi lng ph sa nhiu. S
tch lu cc sn phm ny dn ti s bi lng v hnh thnh cc ng bng chu
th.

9.1.2 p lc do cc iu kin dao ng (nhit , mn, ng vt sn mi)
Vng chu thy triu l vng lun phin ca mi trng sng sng bin v t
lin. Cc thay i to nn s cng thng, kh cn, thiu xy v p lc, v gim iu
kin pht trin.
Vi thy triu rt thp, vng trong triu l mt mi trng khng mang tnh
bin i vi tt c cc sinh vt khng sng trong vng thy triu hoc trong bn ct
su gi nc lnh. Thm ch khi nc c gi li sau mt thi gian di, tnh cht
ca nc thay i ng k. Sau mt trn ma ln, mui ca cc vng thy triu nh
c th gim xung gn nh nc ngt. Vi s gia tng ca cao triu, nhiu vng b
c phi ra. Phn cao nht ca vng trong triu c mang c trng ca mi trng
lc a, nhng sinh vt sng mc triu thp ch b nh hng do b phi ra v tip xc
vi khng kh.
Ti ni triu thp, cc sinh vt bin i mt c vi nng v kh. Trong thi gian
ban ngy b mt nng ln nhanh chng, mc d sinh vt c mt s thch nghi vi nhit
. Nu c th mang nc th tr nn rt nng, nhim v sinh l v thm ch kh nng
n nh ca protein c th b gim. Trong trng hp xu nht, cc sinh vt vng trong
triu c th cht v mt nc. Trong ma h, iu ny xy ra thng xuyn i vi cc
loi yu t.
Nhit v s mt nc thay i hon ton theo phm vi khng gian. C th cho
rng cc ng vt cc vng phng s c kh nng chu nng hn cc loi vng
m v bng rm.
V d ti b bin vng Washington, vo thi k dng thy triu thp trong ma
h bui sng nhit khng kh ch 15
O
C. Ngc li, dng triu thp ma h Puget
Sound li vo gia tra khi nhit khng kh trn 20
O
C v c th ti 25
O
C.
9.1.3 S thch nghi ca cc sinh vt
Kch thc v hnh dng c th nh hng n s mt nhit v nc. Khi kch
thc c th tng th din tch b mt tng tng ng vi khi lng c th, dn ti
gim kh nng mt nc. Do vy, trn quan im mt nc th to s tt hn. Cc ng
vt nh vi b mt tng i ln s mt nhit nhanh hn. S kt hp c hai yu t ny
phi m bo cn bng.

Mt ng vt vng trong triu khng th qu nh, nu khng n s b kh di
mt tri, nhng nu n qu ln, c th b mt nhit nhanh qua b mt ca c th. Hnh
dng cng c nh hng tng t. Sinh vt di v mng s b kh nhiu hn. y l
nguyn nhn ti sao c chn ngng bin c nhng ng nh trong khi triu thp. Vic
thay i hnh dng lm gim din tch b mt v gim s mt nc.

Hnh 9-2: Tn s p ca tua cun (ly thc n) ca mt s loi hu ph thuc vo
s gia tng nhit ti lp trn v lp di ca vng triu (theo Southward, 1964 )

Cc ng vt khng xng sng vng trong triu c th trnh vic qu ti
nhit bng vic lm lnh do bc hi, kt hp vi tun hon cht lng trong c th. Nh
kt qu ca qu trnh , cc loi c sn vng trong triu thng lnh hn cc loi
khc cng kch thc, hnh dng v mu sc. Cc ng vt trong vng triu cao c kh
nng thch nghi tt hn so vi nhng loi vng triu thp v vng triu cao chng
c kinh nghim nhiu gi chu nng hn trong ngy.
Vic di chuyn ca cc ng vt lng gai trong cc loi hu vng thy triu
tng ln vi s gia tng nhit nhng li gim khi gn ti gii hn trn v nhit (hnh
9.2). Cc loi hu vng triu cao chu c nhit cao tt hn nhng loi tm thy
vng triu thp. Cc loi sng trong vng triu cao khu vc nhit i duy tr lng
mao nhit cao nhiu hn cc loi sng vng triu thp hoc triu km.
V iu kin vng triu cao khc nghit hn, nn cc loi y c th chu
nhit v kh. Cc loi sng phn b bin cao duy tr chc nng t bo nhit
cao tt hn nhng loi sng ni bin thp (hnh 9.3). Loi trai m ly thch nghi
tt vi s mt nc v thng chp nhn nhng thay i mnh v nng cu to t bo.
Nhng ng vt sng cc bi bn ct vng triu i mt vi cc vn tng
t ca nhit v mt nc. Ti ni triu thp mc nc h xung, nc tiu thot qua
cc l hng trong bn ct. Trong sut thi gian triu thp cc ng vt c th gi
nguyn trong bn t v lnh bng cch sng trong cc hang tng i su. Nhiu
ng vt khng xng sng bi ct thy triu su 25 50 cm di b mt ct.
Khi triu cao cc ng vt ti b mt hoc qua cc ng ng th. Nhng ni bn

153

ct mn, nc khng tiu thot hon ton, mt s sinh vt nh v mm xp c th sng
gn hn b mt bn ct.
Ti ni triu thp, cc sinh vt bin b phi ra trc tip vi khng kh hoc b gi
trong cc hang hoc vi khi lng nh lng nc gi li. Kt qu l hu ht cc sinh
vt vng triu i mt vi s thiu xy v cc cht thi chuyn ho. Gim tc
chuyn ho l cch thng thng gim xy cn thit trong thi gian triu xung
thp.
Cc ng vt lu ng vng triu c th duy tr c nh ti mt mc triu,
chng tr nn linh hot ti thi gian triu thp, trong khi cc loi khc di chuyn ln
hoc xung theo b bin vi mi chu k triu gi m. Ging nh cc ng vt
bin khc, cc ng vt vng triu thng c tnh thch nghi duy tr v tr. Nhiu
loi c chui cc phn ng c thit k gi v tr tng i vi mc nc.

Hnh 9-3: S tn ti ca cc ng vt c mao trong ba loi ca vng triu v mi
quan h ca nhit vi v tr b bin (Theo Ushakov, 1968.)
9.2 Cc vng ca sng (Estuaries)
Cc ca sng l nhng khu vc khng hon ton kn, ni cc con sng ln
vo bin. Chng c xp vo loi c nng sut cao nht trong cc h sinh thi bin.
Cc ca sng rt giu dinh dng t t lin a ti, nhng nng sut cao ca chng
cn l kt qu ca dinh dng c gi li vng ca sng. iu ny l do lun
chuyn nc khi nc ngt c mt thp hn nm trn nc mn nng hn.

154

Mi ca sng c cc tnh cht vt l ring bit, chng nh hng n sinh thi
ca sng. Nhng tnh cht ny bao gm lu lng dng chy sng, su v a hnh
chung, cc phn lun chuyn c bit, ch kh hu, v thay i thy triu theo chiu
ng. H sinh thi ca sng phc tp hn cc h sinh thi ngoi bin h, v qun x
sinh vt ph du vng ca sng ch l mt trong nhng qun x da trn cc nhm khc
nhau ca cc sinh vt sn xut s cp. Cc thnh phn ch yu i din cho cc ca
sng c m t hnh 9.4. Din tch tng i ca mi qun x ny ph thuc vo
hot ng ca thy triu v a hnh ca sng.
9.2.1 p lc do dao ng mui (fluctuations in salinity)
mui gim dn v pha ca sng c th lm gim s lng loi sinh vt bin.
S bin i ng quan tm nht ca sng l s gim mui khi nc bin vo sng.
Cc loi sinh vt bin c th chu c cc dao ng v mui, nhng cng ch ti
10-25%o. Mt s nhm ng vt khng xng sng (ch yu l ng vt da gai) b
gim vng ca sng do khng c kh nng chng li s gim thp mn. Cc loi
khc nh gip xc c kh nng iu tit tt chng li s thm nn rt ng c
cc vng ca sng. Ti nhng ca sng c s phn tng khi lng ring ca nc theo
chiu ng, mui y ln hn trn mt, dng nc ngt chuyn nc c mui
thp xung h lu. Kt qu l cc ng vt y c th thm nhp xa hn v thng lu
so vi cc sinh vt trn b mt nc. Cc loi sng di nc t kinh nghim thay i
mn hn cc loi trn cn qua mt chu k triu trong ca sng (hnh 9.5).
S chuyn i mn vng ca sng ch yu th hai l khong mui ti hn
3-8 %o. Nhiu nhm sinh vt bin hnh nh kh tm thy s tn ti trong vng c
mui ny, mc d vi nhiu loi c kh nng sng c trong nc ho ton ngt v nc
c mui cao. Trong s cc iu kin thay i ca sng, mt s ln loi d c
tm thy trong khong mui gii hn (hnh 9.6).

Hnh 9-4: M t h thng nhng cng ng sinh vt to nn h sinh thi ca sng (bao
gm mt s loi ng thc vt ch yu) v mt vi tc ng ca con ngi.

155

Cc ng vt thn mm c th khng c kh nng iu tit th tch t bo cc
mui thp. Cc loi sinh vt nc ngt c th iu chnh nng ion v duy tr
trng thi hyperosmotic. Chng mt kh nng iu chnh th tch t bo, v th khng
th chu ng khi mui thp.

Hnh 9-5: S thay i mn trong nc v trong trm tch vng triu ca sng
Pocasset, Massachusetts. Ch rng mn thay i mnh theo su nc do giao
ng thu triu v dng chy sng. Tuy nhin trong trm tch, mn c gi tr trung
bnh v gn nh khng thay i (theo Sanders v nnk., 1965.)

Mc d s thay i mui thng l yu t ti hn trong vic hn ch min
sinh sng ca cc loi sinh vt bin, nhiu loi c kh nng iu chnh v chuyn i
theo s thay i mui. Nhiu loi c c kh nng iu tit rng nn c th vt qua
c s thay i ca mui.
Mt s sinh vt vng ca sng thch nghi chng li vi dng chy ra bin,
nhm li vng ca sng. Dng chy ca sng thng ra bin vi nc c mui
thp trn b mt. Nhng ca sng nh vi thy triu mnh khng th m bo cho cc
loi c non; tnh thch nghi khng chng li s trao i thy triu vi bin m.
Trong phm vi cc h thng ca sng ln nh Vnh Chesapeake, cc nhnh sng nh
sng My c tc trao i thy triu tng i thp vi phn cn li ca ca sng. S
trao i b gim dn n n gi li nhiu u trng ca cc loi ca sng. Ngi ta thy
rng, chng li dng chy ca sng, c vng ca sng v cc u trng/con non ca
cc loi khng xng sng thng tr ng y sng trong sut thi gian triu xung
v chng bi tr li mt nc khi triu ln.
C mi du d b nh bt hn trong thi gian triu ln, chng t rng s thch
nghi ca chng h thng ca sng.

156


Hnh 9-6: S a dng loi ti ca sng Randersfjord, Denmark (theo Remane v
Schlieper, 1971)

9.2.2 Nc l v s a dng cc loi (Brackish water and species diversity)
Nc l l nc cha nhiu mui hn nc ngt nhng li t hn nc bin.
Mi trng nc l cng l mi trng dao ng v bin i. mui c kh nng
thay i theo thy triu, lng nc ngt t sng v hoc t nc ma, v s bc hi
nc. Kt qu l nhiu c nc l c th chu c s thay i mn. Ni chung,
nc l l ni m mui ln hn 0,5%o nhng li nh hn nc bin (35%-37%o).
V kha cnh sinh thi, mi trng bin hng v bin v mi trng nc ngt
hng v t lin to thnh nhng ranh gii khng n nh v nc l to nn vng
chuyn tip. Chng bao gm nc vng ca sng cng nh cc vng m ly ngp
mn, cc vng vnh (nhn nc ngt) v cc m ph ven bin chu nh hng ca
thy triu. Ni n gin, nc l l loi chu nh hng c nc ngt v nc mn. Rt
nhiu phng php phn loi c gng chia cc loi nc l khc nhau; mt s da
vo mui hoc phn vng sinh thi trong khi mt s phng php khc li dng
kh hu, nhng cha c phng php no a cc qu trnh sinh hc hoc sinh thi vo
nghin cu.
V kha cnh sinh hc, tt c cc loi trong cc mi trng ny u thch nghi
vi i sng trong nc c s thay i mn. V mt sinh l, cc nh hng ca ca
mn tc ng ni ti vi nhau v nhit v mc xy ho tan v v th khng th
xem xt mt cch c lp. V vy ti nhng mui nh nhau, s thm nhp vo nc
m s d hn v tng nhit ci thin tim nng iu tit thm lc. C nhiu nh
hng qua li gia cc tham s: v d ion can xi lm tng s khng c i vi mn
thp v n lm gim tnh thm ca mng.

157

S a dng loi trong vng nc l ca sng, mc d cc ca sng c nng
sut kh cao v l ni sinh sng ca nhiu loi c non cng nh c mt s ln cc loi
gip xc, thn mm, chim bin v chim nc, s loi tm thy cc vng ny l tng
i nh so vi cc mi trng sng bin. Mt s t loi c thch nghi ng u
vi mn, nhit v s thay i c trong mi trng nc ny. mn ng
vai tr chnh ca s phn b cc loi vng ca sng. Hnh 8.8 cho thy s phn b in
hnh v tng i ca cc loi ng vt bin, nc ngt, nc l trong mi quan h vi
ca sng. Cc ca sng tr gip cho h ng vt bin in hnh nhng s loi sinh vt
bin suy gim khi nc tr nn t mn v cc loi thay i t iu kin ny sang iu
kin khc. Phn ln cc ng vt sng trong sng khng chu c mn ln hn
0,5%o v chng khng th thm nhp xa hn on ca sng. Ch c mt s t cc sinh
vt nc ngt c th sng trong nc c mui ln hn 0,5%o n 5%o v mt s c
nh c c gai. C mt s t loi c nc l tn ti trong iu kin ca sng vi mui
khong 5-20%o, v hu ht cc ng vt tng thch vi bin. Cc sinh vt bin to
thnh b phn ln cc loi sng ca sng, v s phn b ca chng m rng t bin
ti cc vng trung tm ca sng. Cc loi Stenohalinc khng c kh nng chu mn
thp hn 25-30%o, v chng b loi ra khi cc vng ca sng. Mt s c nh saon hay
ln tm thi sng vng ca sng v di chuyn t do t bin vo sng h, hoc
ngc li. Nhn chung, cc ca sng c t loi hn cc mi trng nc ln cn, nhng
li tp trung nhiu loi c th cng nh sinh khi thng tng ln ng k.

Hnh 9-7: Biu l tng v s phn b s loi sinh vt nc ngt, nc l v nc
mn theo mn
Cc loi nc l cng dc c trng bi a dng. V mt sinh thi, nc l
(vng nc c tng tc vi t lin v hng li t cc ngun vo ca t lin) l
nc giu dinh dng v c gi l nc giu dinh dng. V nguyn nhn ny,
nc l c c trng bi cc qun th t a dang loi. Ngc li, cc loi ny c s
lng ln c th v sinh khi ca chng rt cao. Chng c c tnh ca h sinh thi
giu dinh dng, khng n nh v tr.

158


159
Mt s ln cc loi ch thch nghi tm thi vi nc l. Chng sng mt thi
gian y v sau chuyn ra bin hoc nc ngt sinh sn. Nc l rt c v
cha nhiu cht l lng, cc cht l lng c cc vi khun phn hu. Vi s giu dinh
dng nng sut s cp y rt cao.
9.3 Cc rn san h (Coral reefs)
Cc rn san h l cc cu trc c kh nng chng li tc ng ca sng, do
nhiu loi to nn v thng gn kt li vi nhau.
S pht trin ca cc cu trc ny c s tr gip ca rong vng nu cng sinh
ng vt (RVNCSDV - tn khoa hc l zooxanthellae), to - nhng loi c cuc sng
cng sinh vi san h to rn. San h p v a dng nht trong s cc loi sinh vt
bin. Cc kt cu ln c kh nng chng li sng c tch dn t s pht trin rt
chm ca san h. S pht trin ca cc cu trc ny c tr gip bi to cng sinh vi
san h trong thi gian to rn - c gi l RVNCSDV (loi ny s c m t chi tit
phn sau). To san h, bt bin v cc sinh vt khc, cng vi cc qu trnh gn kt
cng dn ti s pht trin ca rn. Cc sinh vt chim u th l nhng nh lm
khung ca rn v chng to nn khun cho cc rn pht trin. San h v to san h kt
ta Cacbonat canxi, cn cc bt bin to khung cng c th kt ta silic. Hu ht cc
loi sinh vt ny sng theo by n v qu trnh kt ta chm chp y lp b mt
sinh hc ca rn pht trin ln pha trn v v pha bin.
V mt a hnh, cc rn san h kh phc tp. Chng tng i ging vi rng
ma nhit i ch n c nhiu tng, cng c cc vng b ph bng do cc cm san
h pha trn che ph. Chnh v s phc tp ny, c hng nghn loi c v ng vt
khng xng sng sng cng vi cc rn san h - m cho n nay l mi trng sng
phong ph nht trong s cc mi trng sng bin. V d nh cc rn Caribe, ngi
ta tm thy vi trm loi ng vt khng xng sng sng di nhng rn san h gn
kt vi nhau. Cng khng c g l l khi ti mt rn c ti vi trm loi c, 60 loi san
h v vi trm loi c. Trong tt c cc mi trng sng i dng, c l rn san h
l ni c cc mi lin kt sinh hc phc tp pht trin nht.
San h to rn thuc B San h su tia tit ra Carbonat Canxi. San h to rn
(Hermatypic coral) ng gp nhiu nht trong qu trnh pht trin ca rn v c nhiu
rong nu vng ni cng sinh ng vt. Trong khi nhiu chi thuc ngnh c si chm
xut hin cc rn san h, s to rn ch yu l nh mt s chi c si chm thuc
nhm san h su tia - mt nhm ch yu l gm cc loi sng theo qun th. Trong tt
c cc trng hp, cc c th - c gi l polyp (Hnh 9.8), tm thc n bng xc tu
v tit ra phn khung xng carbon canxi. Mt s loi thuc loi cng v bao gm mt
polyp v khung, cn hu ht sng theo qun th v cha hng trm n hng nghn
polyp lin kt vi nhau bi cc m. Bt k l thuc loi no ni trn th cc polyp
cng bt cc ng vt tri ni bng cc thch ty bo trn xc tu ca n v tiu ho vt
mi b nh bng rut m (hay cn gi rut tt). Nh nu s lc, RVNCSDV gp
phn vo qu trnh dinh dng. San h c mt lng ln RVNCSDV v vi ho mc

cao ng gp vai tr quan trng trong vic pht trin ca rn. Nhng san h loi
ny gi l san h to rn (Hermatypic corals) .
San h to rn c th chia lm hai dng: dng khi v dng nhnh, mc d cng
c nhng loi trung gian gia hai loi ny. San h dng khi c hnh dng nh tn ca
n v thng khng ng u. Chng thng pht trin chm, thng khng hn 1 cm
mi nm d bt k mt khng gian tuyn tnh no. Cc qun th san h dng nhnh
thng c hoc dng nhnh cy, hoc dng sng hu. Chng thng pht trin nhanh,
vi tc khong 10 cm/nm. Pht trin tuyn tnh nhanh hn i khi cho php san h
dng nhnh lan rng trn cc rn mt cch nhanh chng. Cc cn bo c khi b gy
san h dng nhnh, nhng cc mnh v li c th bt u nhng qun th mi.

Xc tu
Si tiu
ha
Vch ngn
Vch ngn
Phn nn
Ming
Hng
Khoang v
mch

Hnh 9-8: S ca mt polyp loi san h 6 tia .

9.3.1 S phn b ca cc rn san h.
Cc rn san h chim khong 190 triu km
2
, hoc khong nh hn 1% tng tt
c cc i dng trn th gii. Rn ln nht l Great Barrier tri di dc b bin pha
ng ca Australia vi chiu di hn 2000 km v rng 145 km.
Phn b a l ca cc rn san h, c hai min a sinh ring bit l Thi Bnh
Dng v i Ty Dng. Khi san h v cc loi ng vt c cuc sng gn lin vi
chng tin ho, hu ht cc i dng nhit i trn th gii u ni vi nhau. i Ty
Dng v Thi Bnh Dng ni vi nhau bi ng bin rng Tethys, hay cn gi l
Bin ca ngi Tethy, trc y hon ton kh cn cho n tn cui k Miocene,
khong 10 triu nm trc. Thm ch vo u k Miocene, s phn nhnh bt u xut
hin gia Thi Bnh Dng v i Ty Dng, nhng n cui k ny, ng vt i
Ty dng c s khc bit vi ng vt Thi Bnh Dng, d vn c kt ni gia

160

chng m by gi chnh l Panama, v eo Panama lc khng tn ti. Hn 3 triu
nm trc cha c Eo bin Panama, v nhiu nhm ng vt c h hng vi cc ng
thc vt Thi Bnh Dng sng bin Carib. Sau khi c Eo bin, phn ln cc dng
Thi Bnh Dng sng Carib b tuyt chng. Ni chung, s loi sinhvt cc rn
Thi Bnh Dng gp i so vi i Ty dng, m ln nht l vng Ty Nam
ca TBD (Hnh 9-9). Tng th th c s khc nhau rt ln gia cc loi cc rn Thi
Bnh Dng v TD, v hin nay th c bn l khng c mi lin quan no.
Cc rn san h c th phn thnh san h vng v rn ven b.

Hnh 9-9: Cc ranh gii tng i ca cc rn san h Thi Bnh Dng - n
Dng v i Ty dng nhit i cc nhit bin trung bnh nh nht
9.3.2 Cc yu t lm hn ch s pht trin ca rn
Rn pht trin tt nhng ni c nhit tng i cao, mn ln, nh sng
di do v c thp. Cc rn hon ton nm trong vng bao quanh bi ng ng
nhit 20
o
C v gn nh ch vng nhit i. San h to rn khng th chu c nhit
nh hn 18
o
C v thng chng pht trin tt nht khong 23-29
o
C, d cng c
mt vi loi san h c th chu c nhit ti 40
o
C. V tng th, cc rn san h
thng nm nhng vng bin mn xa khi. Him khi ngi ta tm thy rn cc
ca sng vng nhit i.
Bn cnh nhit , nh sng cng l mt nhn t quan trng i vi kh nng
pht trin ca rn san h chnh v s cng sinh gia san h to rn v RVNCSDV. V
cng nh sng gim theo cp s m theo su, qu trnh pht trin rn dng
di su 25-50 m. Su di mc ny, mt s rn pht trin nh s kt dnh v pht
trin ca bt bin. Tuy nhin, v c bn, s pht trin rn san h l mt qu trnh nhy
cm vi nh sng. C c (s ht trong mt n v th tch) v ph sa u c tc
ng xu n cc san h to rn. Cc khi nc c ngn nh sng v lm gim kh
nng quang hp.

161


162
Cc rn san h sng ni c nng lng ca sng cao, nhng cc con sng
mnh c th ph v cc qun th san h v hn ch s pht trin ca rn. V cc rn
san h cn c nc sch v ph thuc vo a hnh, nn chng thng c xu hng
pht trin cc vng c nng lng sng cao. Nc ng mang cht dinh dng v
ng vt ni ti cho san h, v c th thy r rng l iu c li. Cc cn bo, nh
lc xoy Thi Bnh Dng v bo i Ty Dng, thng lm v cc cm san
h v c th dn ti s hu hoi ca ton rn.
Pht trin rn chnh l s cn bng gia tng trng v n mn sinh hc. Nh
va ni n, cc sinh vt sng rn sn sinh ra carbonat canxi - cht to iu kin cho
cc rn tng trng. Jamaica, mc tng trng l khong 1 cm trong nhiu nghn
nm, c v pha bin v hng ln trn. Kt dnh l qu trnh ho hc lm kt cng cc
rn bng cch phun carbonat canxi vo cc l h, khe nt.
9.3.3 Cu trc ca san h
San h c quan h mt thit vi c chn ngng bin y (c hai u thuc lp
Anthozoa) v c lin quan t hn n sa tri ni, cc dng thu tc di y v thu
tc nc ngt. Khng phi tt c cc loi san h u to rn; mt s l ng vt cng
hoc ng vt qun th c kh nng sng vng nc su v/hoc lnh hn v cc loi
ny c th tm thy khp cc i dng trn tri t. San h to rn dng l cc
ng vt qun th, v mi mt rn c to bi hng t cc c th b nh gi l polyp
(Hnh 9-8). Mi polyp tit ra carbonat canxi bao bc ngoi vi ng knh khong
1-3 mm. Mi polyp c nhiu xc tu cha thch ty bo c th va bt mi va bo
v. Cc polyp c th sn sinh ra mt cm ln hn bng cch phn chia v tnh, hoc
mc mm, v tt c cc polyp trong mt cm c lin kt vi nhau bi cc m ca
chng. San h cng c th sinh sn hu tnh , sn xut ra cc mt tp on u trng sau
phn tn, nh c v to lp nn cc cm mi.
Cc cm san h n l th khc nhau v kch thc, mt s rt ln, c th nng
ti hng trm tn. Dng ca mt cm, nhnh, khi, thu hay np gp, ph thuc vo
loi v c mi trng vt l m san h sng. Cc loi ging nhau c th c dng khc
nhau khi pht trin trong cc vng chu tc ng ca sng khc nhau, hoc su
khc nhau.
9.3.4 Cc rn san h v s a dng loi
Cc rn san h lin tc b tn cng bi s n mn sinh hc. Nhiu loi ng vt
khng xng sng khoan su vo san h. C vt v c ui gai gia mn b mt ca
san h, nhm bin i rng nhng khe nt v o su vo trong thn qun th. Mt
nhm bt bin (h Clionidae) xuyn thng cc khi san h nh vo cc tn cng ho
hc. Mc d s ph hu xy ra trong mt thi gian di, s n mn ny cung cp nhng
ni n nu cho nhiu loi sng rn tn ti v sinh sng. Tc ng chung ca n
mn sinh hc cc rn l to nn hng tn bn ct ht mn m sau c chuyn v
vng nc su hn, i khi ti 250 m hoc hn.


163
mt rn san h c cc k nhiu loi sinh vt sinh sng. Hnh 9-10 ch l s
minh ho cho mt vi loi ng vt chim u th sng rn san h. Ngi ta c tnh
mt rn c th c ti 3000 loi ng vt, v y vn l mt con s khim tn nu nh
ngi ta tnh c cc loi ng vt nh v trung bnh. C th tm thy h sinh thi rn
cc loi i din cho hu ht cc ngnh v lp. Cc rn Thi Bnh Dng - n
Dng l a dng nht v loi san h, vi t nht 500 loi trong ton vng. So vi vng
ny, th cc rn i Ty Dng ngho nn hn nhiu, ch c khong 62 loi san h
to rn. S loi trong cc nhm ng vt gn lin vi cc rn i Ty dng cng
thng thp hn so vi Thi Bnh Dng- n Dng. Ngi ta c tnh c
khong 5000 loi ng vt thn mm TBD, trong khi TD l 1200, v v c c
2000 loi TBD, 600 TD. S khc nhau v a dng loi c th l do s khc nhau
v tui ca cc i dng v thi i a cht tng ng m cc rn c lin quan n.
C l ni c a dng sinh hc cao l nhng vng bin khi nhit i t b nh
hng do cc thay i kh hu. Ty nam Thi Bnh dng c s bin i v thi tit t
nht v c cc rn san h tn ti lin tc nht qua sut cc thi i a cht. Bn c
a cht hc biu din s dao ng ca mc nc bin trong k Pleistocene gy ra s
thay i c bn trong s phn b li t lin v bin ty Thi Bnh Dng v n
Dng. Vo thi k bng h cc i, mc nc bin thp v nhiu vng nm gia
Australia v vng t lin ng nam hu nh kh cn. Vo thi k nc bin cao
nh hin nay, hu ht cc vng ny chm di nc. Thi k mc nc bin thp
c lp cc qun th san h v gp phn vo s hnh thnh nn cc loi khc.
Cc rn san h cng thng l cc bo tng sng.
i i vi cc khu rng gi trong cc qun th sinh hc lu i, cc rn pht trin
mnh phn nh hng nghn nm lch s . Chng l mt phn khng th tch ri trong
cc nn vn ho v di sn t nhin ca con ngi.
Cc rn san h thng c xem nh l t thuc ca th k 21.
Chng ha hn mt kho dc liu khng l trong kh nng cha tr cc bnh
ung th, vim khp, cc bnh ly qua vi khun ngi, virus, v cc bnh khc. V
mi ch c mt triu trong s khong 9 triu cc loi sng rn c xc nh. Gi tr
dc liu ca cc rn san h trong h thng rn Vnh Montego ca Jamaica c tnh
khong t 54 n 85 triu USD.



Hnh 9-10: Mt mi trng sng trong rn san h ngm (minh ha cho mi trng
sng ca h sinh thi a dng ny) :
1 Chim hi u , 2 Sa, 3 C chim, 4 San h dng thu, 5 Roi bin (Mt dng san h sng
nhnh liu), 6 Triggerfish, 7 Qut bin (Mt dng san h sng nhnh liu), 8 C chn ngng
hnh ng, 9 San h , 10 ng vt dng ru, 11San h s , 12 C bm, 13 Ln, 14 Cleaner
fish, 15 San h ng, 16 c, 17 C mang trn (sn bin), 18 Bt bin, 19 ng vt c v, 20
Trai c ln (Tridacna), 21 Pseudochromid fish, 22 Sao bin, 23 San h mm , 24 Cleaner
shrimp, 25 C chn ngng bin, 26 C khoang c, 27 Giun, 28 c (cowry), 29 Qut bin
(nhnh liu).
9.3.5 Tm quan trng ca rn san h i vi vn bo v b bin.
Cc rn san h khng ch c v p k diu v gp phn rt ln vo a dng t
nhin vi nhiu loi c hu, chng cn c tm quan trng cao v mt sinh hc v kinh
t. V san h ly i mt lng ln CO
2
t i dng trong qu trnh vi ho, nn cc
rn ng vai tr quan trng trong cn bng CO
2
ton cu.

164


165
Ngoi vai tr quan trng trong cc lnh vc thuc v sinh hc, cc rn san h
cn c mt tm quan trng trong vic bo v b bin. Chng gp phn bo v b v
nng cao kh nng s dng ca cc bn cng. V khi ng hng khng tr nn r v
thun tin hn, cc rn cng thu ht khch du lch hn v l ngun thc y s
pht trin nn kinh t ca cng ng sng trong vng ln cn.
Cc rn san h cng l vng m gip b bin trnh tc ng ca sng, gp
phn lm gim s ngi cht, mt mt ti sn v xi l. Trn ton th gii, khong
20 quc gia c rt t ngun ti nguyn ngoi rn san h. cc nc ang pht trin,
cc rn san h ng gp 25% cho vic nh bt hi sn, cung cp thc n cho mt t
ngi tnh ring chu .
Rn san h v cc qun x c lin quan nh thc vt bin, cy ngp mn, m
ly l cc ch s kh nhy cm v cht lng nc v s ton vn v mt sinh hc ca
h sinh thi. Chng ch cho php mt bin dao ng tng i nh v nhit ,
mn, c ca nc v cc c tnh cht lng v ho hc khc. Do vy cc rn
chnh l nhng ngi lnh canh tuyt vi cho cht lng mi trng ca chnh chng.
S kim sot chnh xc ca cc rn c th nhn bit c cc thay i trong cht lng
nc hoc cc tc ng do cc hot ng trn b. Cc thay i v kim sot trong
cht lng nc c th gip cc nh qun l ngun a phng hiu c mi quan h
mt thit gia cc hin tng v hot ng xy ra trong vng c lin quan n cc qun
th san h c th no . Nhng mi lin h ny s gp phn pht trin cc k hoch
qun l ph hp cho cc rn san h v cc ngun ti nguyn bin v b bin khc.
Con ngi c mi lin h lu i vi cc rn. Chng l cc vng nh bt c
quan trng, v cng ngy cng chng t c vai tr rt quan trng v kinh t qua vic
thu ht khch du lch. Cc rn gip cho b bin trnh c xi mn v ct khng b
cun tri. Tuy nhin, cc rn nm gn vng b cng cho thy ang ngy cng b e
do v khng c iu kin thun li nh cc rn nm xa cc trung tm hot ng ca
con ngi.
Cc rn san h cc k nhy cm vi s xo trn v hin nay s rn ang b gim
bt. Hi lin hip Bo tn th gii v Chng trnh mi trng lin hp quc (UNEP)
c bo co v tnh trng mt v hu hoi dn ca cc rn 93 trong s 109 quc
gia. Nhiu rn b hu hoi do hot ng ca con ngi hoc do thay i kh hu bin.
S m rng cng ng loi ngi ra gn cc rn thng lm tng cc loi cht
gy nhim cc vng nc gn b. Cc cht ny c th bao gm nc thi ra t
nng nghip, thuc tr su, cc cht gy nhim cng nghip, v nc thi t cc
khch sn v cc sinh hot trn b. Cc hnh thc pht trin ven b dn ti xi mn
t tng khin cho lng bn ct ph ln cc rn tng ln.
9.3.6 Hin tng san h cht trng
San h cht trng l qu trnh lm trng cc qun th san h do khng cn cc
loi RVNCSDV cc m ca polip, khin cho phn xng trng carbonat canxi ca

san h l ra. Thng thng, san h mt khong gn 0.1% RVNCSDV trong qu trnh
thay th v thit lp (Brown v Ogden, 1993). Tuy nhin, nhng thay i bt li trong
mi trng sng ca san h c th lm RVNCSDV mt nhiu hn. C nhng tc ng
hay thay i mi trng c th dn ti s ty trng san h nh bnh, qu nhiu bng,
bc x tia cc tm tng, bi lng nhiu hn, nhim, thay i mn v nhit tng.
San h ch chu c mt s dao ng nhit t 25 n 29
o
C, tu thuc vo
tng ni. San h cht trng l do s thay i nhit trong mt thi gian di. Cc th
nghim cho thy rng san h cht trng xy ra khi nhit nc ln n 32
o
C v duy
tr mc ny (Brown v Ogden, 1993).
Cc th nghim cng ch ra rng tia cc tm tng cng s gy ra hin tng san
h cht trng. Tia UV gy hi cho san h c th tng vng nc tnh. Lng axit
amino ging nh bo t nm trong cc m ca san h gip cho ngi ta xc nh c
mc UV n c th chu c m cha b ty trng. (Gleason, 1993).
C cu chnh xc ca vic san h b ty trng hay nguyn nhn sinh ra hin
tng ty trng vn cha c bit. c nhiu gi thuyt c a ra c gng
lm sng t vn ny. Cc th nghim ch ra rng RVNCSDV n vo rut ca
polyp v sau b tng ra khi polyp qua ng ming (Brown v Ogden, 1993). Tuy
nhin, ngi ta khng quan st thy hin tng ny trong thin nhin. Mt gi thuyt
khc cho rng, san h b tc ng cho to t cht dinh dng hn v v th m to
ri b polyp. To c th sn sinh cc oxit c hi do b tc ng xu v cc cht c
ny c th nh hng n polyp.

Hnh 9-11 : nh mt loi san h cng b ty trng (loi nhnh liu) sng Pohnpei,
Micronesia, chp bi J Hoogesteger. Ch rng khng phi ton b san h b ty
trng, cc polyp bao quanh ra vn khe mnh.

Hin tng b ty trng, hoc b nht mu ca cc ng vt thn mm sng
RVNCSDV xy ra khi (i) mt RVNCSDV gim v/hoc (ii) nng ca cc sc t
quang hp trong RVNCSDV gim (Kleppel v ng nghip 1989). Hu ht cc san h

166


167
to rn thng cha khong 1-5.10
6
RVNCSDV /cm
2
cc m trn b mt sng v 2-
10 pg chlorophyll a trn mt RVNCSDV. Khi san h b trng chng thng mt i 60-
90% RVNCSDV v mi RVNCSDV c th mt 50-80% cc sc t quang hp. (Glynn
1996). Mu sc nht nht ca san h su tia v san h thu tc b ty trng chnh l do
phn khung xng c cht vi ca loi c si chm hin ra qua cc m trong m (gn
nh khng c RVNCSDV b sc t ho).
Nu s ty trng do chu p lc khng qu nghim trng v nu hin tng ny
gim i kp thi, th san h b nh hng thng hi phc li to cng sinh trong vng
vi tun hoc vi thng. Nu hin tng mt RVNCSDV ko di, tc l nu p lc tip
tc v qun th RVNCSDV b ph hu khng hi phc, th cui cng san h s cht.
Ba gi thuyt c t ra gii thch cho c cu ty trng t bo, v tt c
u coi nhit nc bin thay i qu ln nh l mt trong nhiu nguyn nhn.
Nhit cao v cc yu t nh hng n chiu sng l cc nguyn nhn gy s t
gy cc h thng enzym trong RVNCSDV dng bo v chng li cc oxit c hi.
Kh nng quang hp RVNCSDV b suy gim nhit trn 30
o
C, nh hng ny c
t tc ng n qu trnh phn ly ca cng sinh gia to v san h. Nhit gim hay
tng t ngt s lm gim RVNCSDV do khng cn s gn kt gia cc t bo. iu
ny bao gm s tch ri ca t bo ni b si lng vi rong vng nu v s phn tch
tt nhin ca c hai loi t bo.
Cng c gi thit cho rng s ty trng chnh l mt c cu thch nghi cho php
san h pht trin li vi dng RVNCSDV khc, c kh nng chng li tc ng bn
ngoi cao hn. Cc dng RVNCSDV khc nhau tn ti c gia v bn trong cc ch
th san h, v cc dng to khc nhau ni ln s bin i phn ng sinh l hc i vi
nhit v bc x nh sng. S lin kt gia to v san h c th c mt phm vi no
thch nghi trong thi gian sng ca san h. S thch nghi nh vy c th l theo
dng di truyn (gien) hay do ngoi cnh.
Nguyn nhn sinh hc ca hin tng ty trng san h: Do ty trng san h l hin
tng xy ra khi c tc ng xu, nn iu ny c th l do mt hay nhiu nguyn
nhn kt hp. Do , rt kh xc nh mt cch r rng nguyn nhn ca cc hin
tng ny. Cc yu t sau c xc nh l cc nguyn nhn gy nn hin tng ty
trng:
1)- Nhit , Cc loi san h sng trong mt bin nhit kh hp v s thay i nhit
nc bin mt cch bt thng c th dn ti s ty trng. Hin tng ny xy ra
trong iu kin nhit gim t ngt trong thi gian ngn (-3 ti -5
o
C trong vng 5-
10 ngy), hay khi c cc cn kh lnh theo ma. Ty trng xy ra thng xuyn hn
trong iu kin nc bin tng ln. Nhit tng 1-2
o
C mt cch t ngt trong vng
5-10 tun trong ma h s gy hin tng ny.
2)- Bc x mt tri, Ty trng trong cc thng ma h, trong thi gian nhit v bc
x ln thng xy ra khng ng u ni c qun th san h cn v phn nh l ra.


168
Ngi ta nghi ng rng bc x mt tri c mt vai tr trong hin tng ty trng san
h. C bc x hot tnh quang hp (PAR, 400-700nm) v bc x tia cc tm (UVR,
280-400nm) u c lin quan n hin tng ny.
3)- L thin, Cc rn san h phng l ra t ngt trong khng kh chng hn nh trong
thi gian triu rng cc i, mc nc bin xung thp c lin quan n hin tng
ENSO hoc do kin to nng ln cng c th gy ty trng. Nhit ln cao hay
xung thp, tng bc x mt tri, nc bin cn, ho long nc bin do ma cng c
vai tr trong hin tng mt RVNCSDV, nhng ng thi cng lm san h cht.
4)- Bi lng, Khng c nhiu v d v hin tng ty trng san h ch lin quan n
bi lng. iu ny c th xy ra nhng ngi ta vn cha c chng minh rng bi
lng khin cho cc loi RVNCSDV b ty trng.
5)-Pha long nc ngt, Ngi ta chng minh c rng nc b pha ngt mt cch
nhanh chng do ma t bo v dng chy mt cng gy ra hin tng ty trng san h.
Ni chung, hin tng ty trng trong trng hp ny tng i him v ch xy ra
quy m nh, nm gn b.
6)- Cc cht dinh dng v c, nng cc cht dinh dng c bn xung quanh
tng (chng hn nh a-mo-ni-ac v ni-t-rat) lm tng mt RVNCSDV 2-3 ln hn
l gy trng san h. Mc d s ph dng khng trc tip lin quan n hin tng mt
RVNCSDV, n c th gy ra cc tc ng tri ngc th cp nh lm gim sc
khng v tng kh nng nhim bnh ca san h.
7)- Cc d thng sinh hc, Hin tng mt RVNCSDV xy ra khi san h trong mi
trng c nng cht gy nhim ho hc (nh ng), thuc dit c, du tng cao.
V nng cc cht d thng sinh hc cao mi c th gy ra mt RVNCSDV, hin
tng ty trng t nhng ngun nh vy thng ch xy ra trong mt vng ring bit
v/hoc mang tnh tm thi.
8)- Dch ng vt, Mm bnh gy ra ty trng th khc vi cc loi ty trng khc. Hu
ht cc bnh ca san h gy ra cht hng lot hoc cht tng m v bong ra nhng
khi m mm, dn ti tr xng trng (khng nn nhm ln vi san h b ty trng).
Ngi ta xc nh c mt s mm bnh l nguyn nhn gy ra m b trong m.
9)- Phm vi v thi gian v khng gian ca hin tng ty trng rn san h. San h
cc h sinh thi rn san h trong tt c cc khu rn chnh u b cht hng loi t
nhng nm 1870. Tn s xut hin v mc xo trn gy hin tng ty trng
tng mt cch t ngt t cui nhng nm 70. c bo co v hn 60 hin tng ty
trng rn san h trong s 105 rn b cht t 1979-1990, so vi ch c 3 trong s 63 rn
cht trong 103 nm trc .
Hu ht cc cc vng c rn san h ln trn th gii (Carib/Ty ca i ty
dng, ng Thi Bnh dng, trung tm v pha ty ca Thi Bnh Dng, n
Dng, Vnh A rp, Hng Hi) u tng b cht v ty trng trong nhng nm 1980.

9.4. m ly ngp mn (Mangrove swamps)
m ly ngp mn l loi hnh thng thy chim 60-75% di b bin nhit i
v cn nhit i. y l nhng rng cy cao v cy bi m r trong bn ct mm mc
vng cao hn mc nc triu. Chng to nn mt h sinh thi bin duy nht c kho
sinh khi thc vt trong khng trung v l ni tr n cho c cc loi sng bin v trn
cn. Cc loi cy (tn khoa hc l euryhaline) to nn qun x chuyn ho ny chu
c mt bin mn rng v c th mc c ni nc hon ton mn v vng ca
sng, nhng chng ch thch hp vi nhng vng b c bo v c sng nh. S phn
b ca rng ngp mn ging nh s phn b ca cc rn san h, nhng tri di xa hn
ti nhng vng cn nhit i. nhiu vng, m ly ngp mn tip gip vi di b
bin c bo v bi cc rn ngn cch.
9.4.1 Sinh vt hc cc rng ngp mn
V mt sinh thi, qun x cy ngp mn c th c chia ra: (a). rng trn nc;
(b) m ly trong vng triu; v (c) mi trng sng bn ngp do thy triu. Cc vng
khc bit ny gip cho nhng kt hp duy nht ca cc loi nh c m t di y.
Rng trn thy triu (above tide forest) c hnh thnh nh tn che thn v l
cc cy ngp nc l mt mi trng cy ci ni c tr ca cc loi t lin.
Cc loi chim, di, thn ln, rn, c sn, cua cc loi, nhn v cc loi cn trng
khc l ton b cc c dn ca rng ngp mn, trong cn trng l a dng v
ng o nht.
Vng cy trong vng chu thy triu (intertidal swamp) thng cung cp thc
n v nhng sinh c nh khc nhau gip cho qun x cc loi sinh vt bin
a dng hn. Mt s sinh vt sng gi vo cc r cy ngp mn, s khc th
sng trong hoc trn mt bn ly.

169


170
Hnh 9-12: Chu trnh pht trin ca cy c
Vng bn triu (subtidal zone) cng c bn ct v bn loi ht mn vi hm
lng hu c cao v cc vng ct c th l ni rt tt. R cy ngp mn vng
bn triu gip cho h ng, thc vt giu c v to, bt bin, cc loi c v, c
chn ngng, thy tc,... s lng ng o ca nhng loi ny chng t s cnh
tranh v khng gian y kh cng thng.
Cc sinh vt sn xut s cp trong h thng ny khng nhng gm bn thn cc
cy ngp mn m cn c cc loi to y, c bin v thc vt ph du.

9.4.2 Kh nng chu mn
Cy ngp mn ng trong bn trn nhng chic r trng nh nhng cy c kheo
vn ln trn mt nc c mn cao. m r ny c th chia thnh ba dng:
r gc xo ra nh nhng si dy neo chng li nhng con sng mnh v tc
ng do triu.
cc r nh mc ra t cc r gc ly thc n v sng nh ngun dinh dng
phong ph ngay di lp bn.
loi r th ba dng hp th xy nui cy.
B r ca cy ngp mn chnh l h thng lc dng chy chy ra t trong t
lin v o. Nc chy ra bin c lc bn ct y. Nu khng, bn ct c th lp
v lm cht cc qun th rn san h. m ly ngp mn thng ngp trong nc bin
mn, nhng cy th khng bao gi b ngp hon ton. Trong k triu cng, mt t b
ngp nc v cc ht t cng c th b sng cun i.
C ba loi ngp mn. Ngp mn l loi chu nc tt nht, mc kho trong
bn, v thng d nhn bit nh b r trng ging nhng cy c kheo cong queo. Ngp
mn en thng mc cao m khi triu ln th b ngp, cn khi triu xung th l
ra. Chng c c im l b r (r kh) m thng ra t bn bn di tn cy v trng
ging nh mng ty. Cy ngp mn trng khng c dng r c bit v thng sng
ni khng c nhiu nc.
m ly ngp mn c hng nghn con lch vng ca sng chy qua v b chia
ct bi cc vnh v eo bin nhng nhn t khin cho chng tr thnh ni c kh nng
pht trin sinh hc rt cao. y l ni c tr ca nhiu sinh vt, t cc sinh vt siu nh
cho ti u trng, tm, cua v hng trm loi c v chim bin. i vi nhiu loi sinh
vt nh hn, r cy ngp mn va l nh va l ni cung cp thc n cho chng. m
ly ngp mn c bit ti nh l ni c tr ca rt nhiu loi chim him v ang gp
nguy c tuyt chng. Chng che ch cho cc loi b nng trng v nu, mt s loi
dic, chim g kin v i khi ngi ta cn thy chim hng hc hoang d y.
Bn cnh vic cung cp mt h sinh thi rt c gi tr cho cc sinh vt, m ly
ngp mn cn c hai mc ch quan trng khc. Chng bo v t lin, lm gim tc


171
ng do bo bin v bo nhit i gy ra, v nng cao b bin trc s gia tng mc
nc bin thng qua qu trnh to thnh than bn.
9.4.3 Phn b ca cy ngp mn
Cc m ly cy ngp mn thng xut hin cc vng ven bin nhit i ni
chu nhiu nh hng ca cc dao ng thy triu. V mt sinh thi, chng pht trin
tt nht cc vng ca sng, sau l cc doi ct v ngoi cc o hoc cc vnh
c bo v tt vi nc nng v cc bi bn. t ca cc vng cy ngp mn thng
l bn, mt c tnh thng thy ca cc vng ca sng. Trong khi mt s loi cy
ngp mn c th tm thy vng nc ngt, chng pht trin tt nht cc mi trng
nc l.
Cc khu rng ngp mn phn b cc bin nhit i ca th gii. Chng ch c
th pht trin trn cc vng b c che chn khi sng bin; mt khc ht ging s
khng bao gi c th cm xung t hon ho v mc r. Cc b bin ny c th nm
dc theo pha khut gi ca cc o hoc cc dy o c cc rn san h bo v.
Chng pht trin tt cc vng ca sng nhit i ni m chng c th m rng ti a
din tch rng.
Cy ngp mn xut hin trn mt vng a l rng ln hn l cc rn san h v
c th tm thy chng cc ni ngoi khu vc nhit i nh dc theo b bin pha ty
ca Trung, Nam hay Bc chu M, chu Phi, ni m khng c cc rn san h v xa
hn c th thy cc o pha bc New Zealand v gn ca sng Rio de la Plata ca
Achentina. Cc rng ngp mn ny c th thm nhp mt khong cch kh xa v
thng lu dc cc b sng (n 30 km dc sng Fly ca New Guinea, theo McNae,
1968). Cy ngp mn thng khng mc nhng vng cung san h nhiu nng vi
nhng o cao nh Hawaii.
Cng nh cc rn san h, s a dng ca rng ngp mn thay i theo iu kin
a l. S lng ln nht v loi l khu vc Indonesia-Australia-New Guinea; pha
ng Thi Bnh Dng, v phn nhit i ca i Ty Dng li rt t loi (xem s liu
bng 9.1).
Bng 9-1: a dng loi ca cc rng ngp mn cc khu vc a l khc nhau
Khu vc S ging S loi
Australia/New Guinea 16 35
Asia/Indonesia 17 39
Esat Africa/Madagascar 8 9
West Africa 3 5
Weast Atlantic/Caribean 3 6
Eastern Pacific 4 7
(Ngun: Species Divercity, Vol I Ecological Communities by Riclefs and Schluter, p 218 Copyright
1993 University of Chicago Press. Reprinted permission.)



172
Cc m ly cy ngp mn c tm thy dc cc b bin nhit i c hai bn
cu (hai pha ca xch o). Chng c t tn theo cy ngp mn mc . Cc m
ly ngp mn (trong con mt ca hu ht c dn) ging nh vng m ly y mui,
rn v nhn. Qu thc chng l mt qun x rng nh mt cu ni gia t lin v
bin. Cy ngp mn tm thy dc theo cc ca sng nhiu bn ca cc sng ln; v
vng ven bin chu thy triu c che chn nh cc m ph, vnh, lch v ca vo
ca thy triu.
9.4.4 Tm quan trng ca cc vng cy ngp mn (bo v b, a dng loi)
Tm quan trng ca cc vng cy ngp mn c bit t lu i. Chng nui
dng cc loi tm v c, chng cung cp cc cht hu c cho chui thc n ven b k
cn v l nhng ngun dinh dng c gi tr rt ln. Kh nng n nh t nhin ca
chng gip bo v chng xi mn b bin, che ch cho cc vng t lin khi tc ng
ln mi khi c bo v sng do thy triu. Cc vng m ly ngp mn bo v t lin
khi b xi mn v nh hng ca gi bo. Chng cng c tc dng chng nhim.
lc t nhin cc cht thi cng nghip v sinh hot ca con ngi. Cc vng m ly
cy ngp mn cng ging nh cc vng ngp nc khc, chng l thnh phn quan
trng ca chu trnh nc, lm gim dng nc qu ln trong thi gian l.
Ni chung m ly cy ngp mn c cc nhim v quan trng nh sau:
chng cung cp thc n, ni sinh sn v nui dng nhiu ng vt bao gm t
nht 220 loi c, 24 loi b st v lng c, 18 loi ng vt c v, v 181 loi
chim. Cnh l ca cy ngp mn cht trong nc b vi khun phn hy tr thnh
ngun thc n cho cc loi ng vt non.
chng bo v b bin khng b ph v do sng. Chng lm n nh ng b
bin, tng cng s pht trin b bin v c vai tr nh nhng tng chn chng
sng bin v bo. Trong trng hp ny, chng gi ng b v bo v chng
xi l b bin.
chng gi bn ct, cc cht l lng trong nc, cc dinh dng v cc cht c
hi thng qua qu trnh lc t nhin ca chng. iu ny ci thin cht lng
nc ca cc sng vng triu chy vo vng cy ngp mn.
chng hot ng nh nhng vt cn gi lm gim sc mnh ca nhng cn gi
c th ph hy v tn ph t lin.
Thut ng cy ngp mn "mangrove" cp ti c cho nhiu loi cy chu mn
c trong vng ngp mn v c din tch trong cc loi cy chu mn tp trung ln.
Cc loi cy ngp mn sinh sn bng vic ra hoa da vo th phn do ong v
cn trng. Sau khi th phn, ht gi li cy m v ny mm, ra r trc khi ri khi
cy m. Sau n c c hi tt t pht trin trong bn ct, bi bn trc khi b thy
triu cun i. Nhiu loi cy ngp mn ri b ht ca chng trong nhng v bc.
Nhng v ny tri ni trong nc v chm ng ti mt v tr v mc r tr thnh
cy ngp mn mi.
Cc m ly cy ngp mn mang li nhiu li ch cho con ngi, bao gm mt
s li ch sau y:
Vng cy ngp mn c bit quan trng cho cng nghip nh bt c, l ngnh
c li nhun cao.


173
Vng cy ngp mn l ngun giu c v cht t, c th khai thc mt cch n
nh v bn vng.
Cc sn phm khc cng c s dng, bao gm khai thc cy ngp mn cho
mc ch y t.
Mt s ng vt rng ngp mn nh kh, v cc loi khc c th cho mc ch
du lch nn cng to ra thu nhp cho vng ngp mn.
Tm quan trng ca cc cng ng a phng
Cc h sinh thi ngp mn c qun l mt cch bn vng v mang tnh
truyn thng bi cc c dn a phng cho mc ch thc n, thuc y t, cht t, v
vt liu xy dng. i vi hng triu ngi bn a c tr ven b bin, rng ngp mn
to s tin tng, ni sinh sng c bn v bn vng cc nn vn ho bn a.
Vng m ngp mn c kh nng bo v, gip gim thiu cc thit hi do cc
trn bo v sng bin. Ti nhng vng rng ngp mn b tn ph, cc vn xi mn
v nhim mn tng ln nghim trng v i khi gy tn tht nhiu v ngi v ti sn
do cc trn bo tn ph. Cy ngp mn cn rt hu ch trong vic x l cc nh hng
xu ti mi trng nh cy hp th cc nit rt v pht pho, v vy n bo v c cht
lng nc vng ven bin.
9.5 bin su (Deep sea)
9.5.1 Hoang mc ca i dng: thc n hn ch v sinh khi thp
Mi trng bin su c quan tm n nh s n nh v ng nht v
nhiu thng s vt l v ho hc. Nhit nc thng thp (- 1 n 4
o
C) v mn
gi mc nh hn 35, hm lng xy cng khng i. Bn ct mm y c hnh
thnh t t lin hoc t lng chm cc sinh vt ph du bao ph hu ht cc y bin
su. Cht nn cng b gii hn ln bi cc ngn ni gia i dng nh ln t y
bin. Ging nh cc cc dng chy mt, dng chy y bin kh chm (thng < 5
cm/s) song d thay i hn. Bo ln c th ko di vi tun bin v lm cho cc dng
y bin tng tc v c th i hng. Lung cht hu c c th thay i theo
ma v v lng ng t tng trn mt bin xung y bin.
Hu ht cc loi ng vt c mt trong mi trng ti vi nhit thp, p sut
cao v ch yu l nn mm. Cuc thm him ca Challenger cho thy c nhng thay
i s phong ph ca cc loi khc nhau v ng vt y theo s gia tng su. Hnh
9.13 da trn cc kt qu ly mu y t vng rnh su Kurile-Kamchatka Bc Thi
Bnh Dng nhng nm 1950. Kt qu cho thy, v d, cc loi bt bin hnh thnh
mt thnh phn ch yu ca sinh khi ca cc loi sinh vt y ln gia su 1000 m
v 2000 m, nhng chng li t v him di su 2500 m. Cc loi c sao l nhng
thnh vin quan trng ca qun x ti su 7000 m, ti su chng khng cn
na. V tng th cc loi da bin chim sinh khi ln cc su hn 4000 m ni
m bn ct tng i giu cc cht hu c.

Nhiu loi c tm thy cc vng su hn 600 m v nhiu loi b gii hn
rnh su c th. Bng 9.2 thng k mt s loi su hn 600 m v t l ca chng.
Cc loi c hu chim ti 75% sinh vt y cc rnh su Thi Bnh dng. Mc
cao ca cc loi c hu cho thy rng cc rnh su l cc mi trng c bn b bit
lp, chng l nhng trung tm sinh ra cc loi mi.
Trong bin su, cc cht l lng c chc nng nui dng lp bn ngho y.
Cc hot ng nui dng v chn vi cc ngun thc n b lng ng lm tng lng
nc ca cht lng ng v chuyn chng thnh nhng ht nh c th b ra tri d
dng bi cc dng chy y.

Hnh 9-13: T l phn trm cc nhm ng vt trong sinh khi ti cc su khc
nhau vng Kurile-Kamchatka

Sinh khi thp, mc d vng bin su c kh nhiu sinh vt song cc qun x
sng y bin bn c bi mt sinh hc v sinh khi thp. S lng c th y trn
mt n v din tch gim theo dng hm m vi s gia tng ca chiu su v khong
cch t b bin. Di cc vng gia cc i dng, tng mt cc sinh vt ln thay
i trong khong 30 n 200 c th trn 1 m
2
.

Sinh khi trung bnh ca cc vng su khc nhau nh bng 9-3. Sinh khi sinh
vt y cao nht ven bin nc nng thuc vng quang hp, v thp nht cc khu

174


175
vc thiu kh nng quang hp. Cn nh rng su trung bnh ca i dng th gii
l 3800 m, hu ht y bin y sinh vt sng ch t di 5 g trng lng t trn m
-2
.
Bng 9-2 Phn trm ca cc loi sng su di 6000 m.
Nhm c hi S lng loi % loi c hu
Foraminiferans
Sponges
Cnidaria
Polychaetes
Echiurid worms
Sipunculid worms
Crustaceans
barnacles
cumaceans
tanaids
isopods
amphipods
Molluscs
aplacophorans
snails
bivalves
Echinoderms
crinoids
holothurians
starfish
brittlestars
Pogonophorans
Fish
128
26
17
42
8
4

3
9
19
68
18

3
16
39

11
28
14
6
26
4
43
88
76
52
62
0

33
100
79
74
83

0
87
85

91
68
57
67
85
75
Bng 9-3 Gi tr sinh khi trung bnh ca cc ng vt y cc su khc nhau
Khong su (m) Sinh khi trung bnh
(g wet weight m
-2
)
Vng triu
n 200
500-1000
1000-1500
1500-2500
2500-4000
4000-5000
5000-7000
7000-9000
>9000
3 x 10
3
200
<40
<25
<20
<5
<2
<0,3
<0,03
<0,01

Sn lng sinh vt y khng th nh gi trc tip t cc gi tr sinh khi
nhng nhiu loi bin su pht trin kh chm v sinh khi nh chng t sn lng
thp. Cc c tnh khc nhau cho thy nng sut s cp hng nm trn hu ht cc y
bin su khong 0,005 0,05 g/cm
2
.
9.5.2 Tnh a dng sinh hc cao
a)- Cc iu kin thc n ngho nn, bin su cc vng y bin thoi v mm th c
nhiu loi hn vng y bin dc
b)- Tnh a dng sinh hc ca cc vng y mm tng ln theo chiu su, nhng sau
gim dn khi t n su gii hn khong 2000 m. C th cc su ln hn
2000 m th ti xp ca y bin nh hng n cc iu kin sinh hc. Quy m v
dn s qu thp nn tnh a dng sinh hc thp xung. Gi thit ny ng vi s suy
gim c bit mnh ca ng vt n tht. Trong cc qun x, cc ng vt n tht
thng t hn v s lng v sinh khi v chng ph thuc vo cc ng vt khc lm
thc n cho chng. l nguyn nhn khi ton b cc thc n b suy gim, mt s
ng vt n tht u tin s bin mt..

Hnh 9-14: S thay i a dng loi ca cc nhm ng vt y theo su nc. S
lng loi c tnh i vi nhng loi sinh vt c t 50 c th tr ln (theo ti liu
ca Rex, 1981)

c)- Mc d s tng t khng r trong tnh cht ca bn ct, c mt s thay i
kh r v s a dng theo v tuyn trong sinh vt y bin su (hnh 9-15).
d)- S a dng sinh hc bin su c th c tng cao nh mi trng n
nh, mc d s xo trn v s kt hp cc mi trng sng cng c a dng gia tng.
S a dng sinh hc cao ca bin su tng l mt ngc nhin ln i vi cc nh

176

sinh thi. Mc d khng c nh sng, ngun cp thc n rt km v hot ng ca cc
sinh vt nh rt thp, mt s lng rt ln cc loi c pht hin.

Hnh 9-15: S a dng loi ng vt chn ging, thn mm, hai mnh v theo v
Atlantic. (s liu c ly t Rex v nnk., 1997.)

9.5.3 Cc dng nc nng v cc dng chy lnh
Dng nc nng l mt ct nc nng phun ln t y bin. N phun ra lin tc
nc cc nng v giu khong cht, rt tt cho cc cng ng sinh vt. Mc d hu ht
cc vng bin su mt sinh vt tha tht song rt nhiu sinh vt sngc bit tp
trung ti nhng ni c dng nc nng. Cc loi giun ng v trai ln thy rt nhiu
cc vng c khe dng nc nng Thi Bnh dng (Pacific), trong khi li loi tm
khng mt ch thy cc khe nc nng Bc bc dng(Atlantic).
Khe dng nc nng u tin c pht hin vo nm 1977. Cng tn ti cc
i dng Pacific v Atlantic. Hu ht c tm thy su trung bnh khong 2100
m nhng vng y bin dc theo h thng Mid-Ocean Ridge. (hnh 9-16).

177


Hnh 9-16 H thng dng nc nng dc theo dy Juan de Fuca. Hy quan st s
chuyn ng ca nc bin qua cc khe nt trong Basalt
Nhiu loi ng vt c mt cao trong cc qun x vng dng nc nng c
mt quy m rt ln. S xut hin sinh khi sinh vt y rt cao nc su, khng dng
cc sn phm quang hp t lp mt, t ra cu hi l lm th no cc ng vt ny
c c thc n.
Phn ng i hi xy c cung cp bi nc bin xung quanh. Tnh cht sinh
ha ca sn phm ha tng hp c th c tm tt nh sau:

CO
2
+ H
2
S + O
2
+ H
2
O CH
2
O + H
2
SO
4
(9-1)
carbohydrate
Trong cc cng ng sinh vt ca dng nc nng, vi khun tng hp ha l
sinh vt sn xut s cp ca chui thc n, v sinh khi ca chng tr nn c sn cho
cc ng vt bc cao hn. Nhng mng vi khun dng tc to nn thc n cho cc loi
ng vt khc.
Khong 95% cc ng vt tm thy vng dng nc nng l nhng loi
c bit t trc. n nay gn 300 loi mi c m t, nhiu yu cu thit lp s
phn loi mi v chng qu khc nhau gia cc loi lin quan.
Cc dng chy lnh l nhng vng giu cht lng metan v sulfit chy vo nn
y bin. Ging nh cc loi vng cc dng nc nng, cc ng vt dng nc
lnh cng ph thuc vo xy ha sulfur ca cc vi khun cng sinh tn ti. Cc
dng nc lnh l nhng vng tng t nh cc vng ca dng nc nng, mc d

178


179
nc y lnh vi cng nhit nh nc xung quanh, chng c gi l cc dng
nc lnh l so vi nc cc nng ca cc dng nc nng. Cc dng nc lnh tr
gip cho sinh vt sng tng t nh trong dng nc nng thng qua s pht trin ca
cng ng sinh vt xung quan, chng ph thuc vo cc ha cht nh l hydro sulfit,
metan, st, mangan v silic c trong cht lng vng nc lnh.
Kt qu dng chy lnh do cc nguyn nhn khc nhau, v chng thng thy
dc theo cc ra lc a v trong cc vng m lp v mt i dng c chm xung
lp di ca v tri t. Cc qun x vng nc lnh cn ph thuc vo sn phm ha
tng hp ca cc vi khun sulphide m khng phi l quang hp.
Vic pht hin cc dng chy lnh di i dng cho thy rng thnh phn
quan trng nht cn cho nng sut sinh hc cao trong bin su l mt ngun cc hn
hp hu c suy gim khng nng. S tn ti cc dng nc nng v dng chy lnh
trong nc bin su ch ra rng nhit thp v p sut cao khng lm gii hn cc
hot ng ca sinh vt sng bin su. Trong cc vng nc su, ni m sinh thi sinh
vt y ph thuc vo sn phm quang hp ti tng mt, cc qu trnh sinh hc v sn
phm y b gii hn v rt t thc n. Trong nhng vng chui thc n da vo
sulphide, nng sut sinh hc trong bin su c th vt qu vng ph dng
(euphotic).
Mc du mi dng nc nng hoc dng chy lnh c cc tnh cht vt l khc
nhau v h ng vt, cc qun x ng vt kt hp vi nng sulfide cao tng t
nhau trong mt s kha cnh. Cc loi quan trng nht thng tng t v sinh thi,
nu khng lin quan n phn loi. Tt c cc cng ng sinh vt c c trng bi
mt dn c cao, sinh khi ln, v tc pht trin nhanh.
y l nhng s tp trung duy nht ca i sng trong nhng vng bin su m
thng c trng bi mt v nng sut thp; cng nh vy, cc cng ng sinh vt
sulphide ny c hiu mt cch chnh xc nh l nhng "c o".
Tng hp ha bi vi khun l ngun thc n c bn trong tt c cc cng ng
ny. Chng cn nh sng v sn phm quang hp t b mt bin a xung; dn c ca
sinh vt sng hon ton n nh nh cc cht v c c chuyn i sang sinh khi vi
khun, ci m sau tr nn c sn cho cc ng vt cao hn tiu th.
Nh vi khun, cc loi giun ng v trai ca cc dng chy lnh pht trin khng
cn nh sng mt tri, chng l loi ging nh cy i vi h sinh thi bin su. Thc
n ca chng l nhiu loi dng vt khc, v y l mt trong nhng lnh vc chng ta
nghin cu.
Cho n nay, rt t hoc khng c thng tin sn lin quan n qu trnh sinh thi
nh hng n tc sinh sn ca qun c cc vng dng chy lnh di bin su.
n tht ln nhau, cnh tranh v s ri lon ng vai tr c yu, nhng t cc gi thit,
nghin cu lin quan n qu trnh pht trin dn c ca cc vng nc ny.
9.5.5 C bin (Sea Grass)
C bin mc khu vc ca cc m ly nc mn, trong vng ngp triu hoc
nh hng thy triu v n c th c cc loi c di trong . Ni chung, c di khng
mc tt trong nc vng ca sng c bn nh chng thng mc nhng vng nc
trong. Loi cy chim u th ca qun c vng ca sng nhng vng n i l


180
Zostera, thng c gi l c ui ln; cc vng kh hu nhit i, n c thay
th bi c Thalassia hay c b st y. C di mu nu Fucus v cc loi c di xanh
c th mc cc vng dc cc vng y c nhiu c bin mc.
Mt s ln h thc vt nh bao gm ng vt nguyn sinh (n bo) v mt s
loi khc kt hp vi cc loi c bin v cc ng vt khng xng sng ly thc n
qua b phn lc tn cng vo l ca c bin. Cc loi c, ng vt hai mnh v (trai,
ngao,...), v cc loi ng vt v cng khc (tm, cua) chim u th thng tr h ng
vt khng xng sng di chuyn trong cng ng c bin. Hn na, vng ca sng l
vng nui dng con non ca rt nhiu loi c, k c nhng loi c c gi tr thng
mi rt cao nh c hi, ...
C bin l loi cy c hoa, nhng hoa li rt dn gin v phn hoa c nc di
chuyn d dng. Chng xut hin cc vng nc nng nhit i, cn nhit i v c
n i. Cc loi c bin nh Pbyllospadix ca Thi Bnh dng c th mc vng
nc c sng nhng cc loi khc li mc vng chu nh hng triu c y mm
kh yn tnh v nng. su ln nht m chng c th mc chu nh hng ca nh
snh, v th c c th lm gim ng k n su m chng c th mc. Loi c
ui ln mc (Zostera) rt nhiu nhng vng y bin nng ca Bc Bng dng,
pha ng Thi Bnh dng, v a trung hi. Cc loi c khc c th tm thy khp
cc vng ven bin trn th gii.
C bin thng m rng qun c ca chng bng pht trin v tnh nh l h
thng thn r st mt. Cng c nhiu loi c bin sinh sn hu tnh v ht ging biu
th mt phn s gia tng s lng. S lng c bin thng m rng ch yu bng vic
m rng thn r ca chng. Dinh dng c nhn t nc qua cc l nh ca lp nn
y bin, mc d mt s c bin c cc vi khun c nh m cng sinh trong cc h
thng r ca chng, nhng vi khun ny cung cp nit.
Cc y bin c c d dng chim c bn ct lng ng sau ginh vai tr ni bt
ni tip lin tc s chim ng trc ca c di. Mt vng y c bin pht trin tt
c th m rng sau ti cc bi trng ph sa lng ng bng cch m rng h thng
thn r. Bi ph sa trng, thng c t dinh dng. H thng thn r tr gip trong vic
chuyn dinh dng t mt bin c hnh thnh. S chim c ca cc vng mi
bng ht l kh khn tr khi ph sa n nh v giu cc cht dinh dng b phn
hy. iu ny c th c t c bng s c mt ca cc thc vt khc, v d c di
chng lm n nh ph sa lng ng v b sung dinh dng trong cc dng nh
amomonium. V vy, bi ct trng c th thay i bng s ni tip nhau c mt y
c bin (xem hnh 9-17).
Cc vng y c bin c nng sut s cp cao v gip cho hng lot cc nhm
loi ng vt sng trong pht trin.



Hnh 9-17: S chuyn ho y bin ct sang y bin c Thalassia vng Florida
9.6 Cu hi v tho lun

181


Chng 10
Tc ng ca con ngi

10.1. Tc ng ca con ngi n cc h sinh thi nc
Cc h sinh thi chu s tc ng ca loi ngi (human impacts) nhiu mt
(Hnh 10-1). Nhng tc ng ny c th trc tip, v d nh xy dng p dng, thi
nc bn, khai thc qu mc ti nguyn thin nhin ( c trong t lin v ngoi bin),
hoc c th l gin tip nh nh hng ca s thay i kh hu hay s dng t. Phn
ny s a ra mt ci nhn tng quan v mt vi tc ng gy ra do cc hot ng ca
con ngi ln vng ca sng, vng ven bin v ngoi bin.

Dng chy
mt
Cng
nghip c
Cng trnh thu li
Cc cht
c
Kh
hu
Ma
axit
Ph dng

Hnh 10-1: Cc tc ng ln thu h sinh thi
Trong s cc tc ng gy ra do hot ng ca con ngi, cng trnh thu li,
dng chy mt v cc cht thi c hi t cng nghip v nng nghip... l cc tc
ng trc tip n h sinh thi nc (sng, h v bin). Ph dng, ma axit v s thay
i thi tit thng l cc tc ng gin tip.
182

Trng i hc Thy li H Ni- Vin nghin cu thy lc Deltf 183
10.2. Tc ng ca con ngi n vng ca sng v vng
ven bin
Rt nhiu thnh ph ln nht trn th gii nm vng ca sng, chng hn nh
Lun-n, Thng Hi, Niu-oc... iu ny khng c g ng ngc nhin, do vy m
nhng vng ca sng ny chu nhiu tc ng do con ngi gy ra hn bt k mt mi
trng bin no. Vng b nh hng nghim trng nht thng l vng c ni m ly,
v vng ny thng c s dng xy dng v t cc khu cng nghip, v i khi
cn c lm sn bay. mt vi vng ca sng, hn 90% bi c m ly bin mt.
Bn ct ven bi vng nh hng triu v cc loi thc vt bin thng c no vt
lm cng ln hoc cng trnh nh hn nh bn du thuyn.
Cc c im to iu kin thun li cho cc vng ca sng m mang pht trin
cng chnh l cc c im khin cho nhng ni ny tr nn d b nhim. Cng nh
cc cht dinh dng, cc cht gy nhim nh sn phm ph t cng nghip ho du,
kim loi nng, phn bn v thuc tr su c gi li trong h thng. Cc cht ny
khin cho t tr nn ph dng mt cch khng mong mun v sinh ra nhim, v
cc sinh vt gy bnh, kim loi nng v thuc tr su ny c th tn cng vo dy
chuyn thc n thm nhp vo thc n ca ngi.
V vng ca sng l vng c iu kin t nhin thun li cho pht trin, cc
vng ny cng thng c khai thc lm ni nui trng thu hi sn v cc loi thu
sn c v. Tuy nhin, gn y, cc khu cng nghip thu sn vng ca sng v khu
nui cc loi tm cua b ng ca do mt vi khun coliform cao thi ra t cht
thi sinh hot, kim loi nng trong cc sn phm hay do s tch ly thuc tr su. Tt
c cc dng nhim ny cng lm bin i cu trc ca h sinh thi ca sng, thm
ch cc vng nhim nng cn thiu xi trm trng m ch c vi khun mi c th
sng c.
Nhn chung, lch s ca s tn ti cng nh pht trin vng ven bin khp tri
t cho thy s suy gim ngun ti nguyn ven bin v mt dn s a dng sinh hc.
Cc mi trng sng c bn b ph hu, cc qu trnh trong h sinh thi b gin on
v nc b nhim mt cch trm trng. Tt c cc tc ng ny u do con ngi gy
ra v c gi l cc tc ng do pht trin (development impacts). Nh chng ta
bit, vng ven b bin l ni con ngi sinh sng, cy cy, bun bn, xy dng cc khu
cng nghip, gii tr v cng l c s cho cc hot ng thuc lnh vc hng hi nh
vn chuyn bng tu thuyn, nh bt hi sn v khai thc cc ti nguyn bin. S s
dng ny khng phi lc no cng tng thch nhau m c th mu thun nhau trong
vic s dng v ra quyt nh. Cc vn sau y l cc tc ng do pht trin chnh
ngay vng b bin hoc thm ch c vng xa b.
1)- Nng nghip (Agriculture), cng nghip ho nn nng nghip vi vic tng cng
s dng cc ho cht e do mi trng vng ven bin. S dng phn bn ngy
cng tng l nguyn nhn chnh gy hin tng ph dng trong sng v vng ven bin
do nitrat d tha ngm vo nc mt v nc ngm. Cc sn phm cng nghip

(chng hn nh thuc tr su, thuc dit c v phn bn) c s dng ngy cng
nhiu trong nng nghip, trong khi nn sn xut nng nghip ang s dng nhiu ho
cht trong cc qu trnh ch bin. Thuc tr su c th git cht hay lm suy yu cc
sinh vt sng bin khi ch cn c mt trong nc vi mt nng nh.
2)- Thu sn (Aquaculture), Cc vng nui tm v khu nui trng hi sn thng nm
ranh gii gia bin v t lin, t hu dng (h nui tm), vng t t (tambak) v
bin (lng hay cc mm nhn to nui c hi). Ngnh thu sn l mt trong nhng
ngnh pht trin nhanh nht trong nn kinh t vng ven bin. Tuy nhin, nu s m
rng ngnh ny khng c kim sot, hu qu lu di c th l lm gim gi tr ngun
ti nguyn v nhim vng nc nng ven bin.
3)- Cng nghip rng (Forest industries), Trong cc lu vc, din tch rng ang mt
dn lm tng bn ct trong sng. Cht lng nc gim v c tng. Dng chy mt
tng c th cun v bin cc cht nhim ho hc di nhiu dng khc nhau. Mt
nh hng nghim trng nht l lm ngt th cc va san h v thm ph thc vt
bn ngp (c bin v rong bin) - cc yu t quan trng trong mi trng sng ca c.
Dng chy rn cng lm cho cc loi ng vt c v nh trai, s cht v lm gim sc
sng ca cc ng vt vng ven bin cng nh rng c mn.
4- Cng nghip nng (Heavy industries), Cc khu cng nghip nng e do n c
nhng vng nm cch xa v tr nh my. Nc thi t cc khu cng nghip nng ven
bin c th gy nguy him nghim trng ti cc h sinh thi ven bin. Cc nh hng
ny c th l nhng xo trn nh (nh tng c cc b v tm thi) hay nhng bin
i rt ln (nh nhim nc do cc cht ho hc c hi thi ra).
5)- C s h tng (Infrastructures), c s h tng pht trin, nh in, nc, gas, cng
rnh, ng s, ... c thit k phc v cho nhiu ngi s dng v thng c
ln k hoch v xy dng bi Chnh ph. Cc h thng c s h tng pht trin vng
ven bin c th c nhiu loi tc ng n mi trng (Hnh 1-8). Bn cht ca cc h
thng c s h tng l tuyn tnh. Cng nh vy, mt vi h thng c th l ng kt
hp, ng dy, ng ng cung cp cc hnh lang pht trin tuyn tnh hay cc ht
nhn thng mi.
6)- Khai thc m (Mining), ct xy dng l mt mt hng c gi tr, nhng cn phi
nh rng yu t c bn trong vic bo v thin nhin cho khu ven sng, bin chnh l
ct. Ct khai thc c gi li cc kho ven b v sau n lm tc nhn cn lc tc
ng ca sng bo.
7)- An ninh quc phng (National security), Vng bin v ven bin l ni c dnh
mi quan tm c bit v vn an ninh v y l vng ranh gii, ni c th xy ra s
xm lc v cc hot ng tri php (nh bun lu). Cng bin v bn tu, cc sn bay
ven bin v nhiu c s c bit cho nhiu lnh vc c t vng ny, v vy chng
c u tin m bo v an ninh.

8)- Cng nghip ho du (Petrolium industry), Cc cht thi t du l mt mi nguy
him thng trc i vi cc h sinh thi bin, nhng ch c mt s t quc gia c k
hoch phng b v i ph trong trng hp khn cp mt cch y . Hu qu ca
vic du trn ra bin cng ngy cng tr nn thy r nhng c l khng nghim trng
bng nh hng ca vic thi cc cht hu c v ho hc c hi.
9)- Cng bin v bn du thuyn (Ports and marinas), cng v cc phng tin h tr
cho vn chuyn hng hi thng phc v cho 4 hot ng chnh sau: (a) khai thc du
kh ngoi khi ; (b) cng nghip vn ti hng hi; (c) khai thc hi sn; (d) cc hot
ng v qun s. Cc e do chnh t pht trin cng bin l vic chim vng t t
ni c rng c mn lm bi xy dng, thi t t no vt knh, cc hot ng
o p v ph hu cc bi c ven bin, x cc cht nhim ra bin.
10)- Du lch (Tourism), suy gim mi trng e do n cng nghip du lch qua nhiu
cch. Gim a dng sinh hc, ti nguyn cn kit, v cc vn v sc kho ca con
ngi c th l do nh hng v mi trng c tch ly dn t du lch. Mt tc ng
nghim trng t vic pht trin du lch trn khp th gii l cht lng nc gim do
nc thi gy ra.
11)- nh c (Settlements), con ngi c th tp trung nh c ng vng ven bin
vi nhiu loi tu thuyn, khai thc ti nguyn qu mc, lm nhim t vic xy dng
cc khu cng nghip, thi mt khi lng ln rc v cc cht thi ho hc. Nhng vng
t ven bin l thc s cn thit cho con ngi nh c, nng nghip, bun bn, cng
nghip, gii tr, v cc hot ng h tr cho vn ti hng hi cng nh nh bt hi sn.
12)- Cc cng trnh bo v b (Shore protection works), Cc gii php cng trnh
chng xi mn b v bo v cc c s h tng ven bin khi nguy c b tc ng do
sng bin rt tn km v thng mang tnh tm thi hoc phn tc dng. K, bin,
chn sng, v cc cng trnh bo v thng dng khc thng c tc ng ph phc
tp v khng on trc c, gy ra xi l pha h lu, v c khi lm cho b bin mt
hon ton. Cc mi trng sng vng bin, nh cc loi c, cc va san h, cc o
nm ven bin, h mn, thng c cc chuyn gia nghin cu v him ho thin
nhin xem nh cc tc nhn bo v tt nht chng li bo bin v xi l, lm lch
hng v tiu tan phn ln nng lng ca bo bin. Do , vic duy tr cc mi trng
sng t nhin ny l rt quan trng trong vic bo v b cng nh bo tn mi trng.
13)- Cht thi (Waste disposal), Cc khu dn c ven bin thi ra mt lng ln cht
thi. Mt vi ni c nhng b cha cht thi r r vo tng nc nng. Mt s ni tp
trung cht thi li, x l ri thi ra bin. Cc bn cng cc thnh ph ln ven bin v
cc cng cng nghip thng gp vn v thi cc cht gy nhim. Hai loi
nhim chnh cc cht thi hu c tiu th xi v cc cht thi cng nghip c hi
ph hu mi trng v ti nguyn vng bin v gy ra nguy him cho sc kho con
ngi. Mt vn khc na l lng ln cht thi rn c chn dc theo b bin lm
hng cnh quan thin nhin v lm cht gy nhim ngm vo vng bin ven b.
14)- Cc d n cung cp nc (Water supply projects), sng chuyn cc cht dinh
dng c gi tr vo cc h sinh thi bin v to nn mi trng nc l thun li cho
cc rng c v c ging m vng ca sng, cng nh l ni sinh sng ca cc

loi chim chuyn sng ven vng m t v cc loi khc nh c su. p dng v cc
cng trnh phn dng hay ly nc c th lm mt cn bng nghim trng cc h sinh
thi c ngun ti nguyn ph thuc vo sng v lm gim s pht trin v a dng sinh
hc do chuyn nc t h sinh thi hoc do s dng cc bin php iu tit nhm
thay i chu trnh dng chy phc v cho ti tiu, kim sot l, cung cp nc, ...

Sinh thi
hc
Mi quan
tm ca
con ngi
Ho-l
hc
Mi trng nc
Mi trng cn
Cng nghip c *
Ph dng *
Thc vt sng di nc *
a dng loi *
Cc loi gp nguy him *
*Rng
*Sinh vt hoang d
*a dng loi
*Cc loi ang gp nguy
him
t Sc kho
Xi v bi *
nh hng nc vt *
n nh ca b *
Tiu thot nc *
c tnh ca t*
* Bnh tt
* V sinh
* Dinh dng
Nc mt
Kinh t - x hi
Nc ngm
Khng kh
Cnh quan
* Mt t
* Sn xut ma v
* Thu sn
* Ti tiu
* Giao thng thu
* Kim sot l
* Giao thng
* nh c
* Lao ng
* Cng-nng nghip
* Ch thu vn theo vng
* Ti trng bn ct
* nhim nc
* Ch thu vn theo vng
* Ngm vo tngngm
* Mc nc ngm
* nhim nc
* nhim khng kh
* nhim do bi
* nhim do ting n
* Phong cnh
* Khu vui chi gii tr

Hnh 10-2: Cc tc ng "di nc" (coastal waters) and "trn cn cn c kt
hp xem xt khi nh gi nh hng ca c s h tng n mi trng vng ven bin
186

15)- Khai thc bin (Marine excavation), vic khai thc bin, hoc no vt s lm
xo trn hay lm bin mt cc sinh vt sng y bin, gy bn ct l lng trong
nc, tch ng chng mt ni no di y, lm gim kh nng xuyn sut ca
nh sng, tng c, thay i s lu thng, gim xi ho tan, tng dinh dng
trong nc. S bin mt mi trng sng ca cc sinh vt y trong qu trnh no vt l
kh trnh c nhng s ph hu cc vng khng b no vt nm k , nh va san
h, l c th trnh c. Nhiu h sinh thi vng bin (chng hn nh cc va san h)
rt nhy cm vi bn ct l lng v bn ct di y, v cn thi gian di pht trin
tr li. Trn thc t, gim tc ng do no vt l mt vn quan tm chnh trong khi
nghin cu v k thut v mi trng.
16)- nh bt thu hi sn (Fisheries), nh bt hi sn qu mc gy ra s xo trn
kh ln trong cu trc dinh dng ca tng nc. Hn na, cc loi nh ca li
thc n thng c kch thc c th ln v chng l nhng sinh vt n tht rt linh ng
bt v n tht cc con mi ln. Vi sinh khi thp v kch thc ln, cc sinh vt n
tht lp trn thng khng c nhiu. Chng ta s thy r hn nu nhn vo s s t
ni vi s hu nai hay s chut. Cng tng t nh vy, nhng sinh vt n tht lp
trn l cc loi c a thch ca con ngi, bao gm cc loi c ng, c hi,...
Nu nh bt hi sn tp trung vo nhng loi lp trn tng i him ny, chng ta
nn d on n s bin mt ca ngnh cng nghip nh bt hi sn vo mt ngy
khng xa. Daniel Pauly v ng nghip nhn ra iu ny qua mt phn tch ton cu
v cu trc dinh dng cho ngnh thu hi sn. H s dng mt b ti liu thu thp
nhiu ni trn th gii v cng nghip c c T chc Nng nghip v thc phm
(FAO) ca Lin hip quc bin son li. Trong 45 nm qua, nh bt c chuyn t
loi c ln n c sang loi c n ph du v ng vt thn mm. Mt khc, nh bt cn
kit cc loi n tht lp trn c th lm tng cc loi mi, v do vy m c th lm tng
ln ca n mi. Nhng suy ngh ny vn cha y . u tin, r rng l p lc
nh bt loi n c (c ng v c hi) l khng th trnh c. Th hai, chng ta thch
n cc loi ny. Th ba, khng c l do g tin rng chng ta s khng nh bt qu
mc cc loi cc bc dinh dng thp hn, khi dn s th gii ngy mt tng nh vy.
10.3. nh hng ca cc hot ng con ngi n h
sinh thi bin
Con ngi tc ng n h sinh thi bin ch yu do gy nhim v khai thc
qu mc ngun ti nguyn bin. Phn ny s m t mt cch s lc v s nhim v
cc hot ng khai thc ca con ngi.
10.3.1 nhim bin (Marine Pollution)
Ni chung con ngi gy ra nhim, tc ng n mi trng bin cc
kha cnh sau:
a)- Cc tc ng phc tp ca con ngi lm cho vic xc nh vai tr ca cht
nhim trong s suy thoi mi trng bin tr nn kh khn hn. Cc nh hng ca
con ngi n mi trng bin c th c phn ra lm cc loi c bn sau:

1. S thay i tng nn do b no vt, thay i cu trc b v do bi lng.
2. S xut hin cc cht c nguy him i vi c s sng ca sinh vt bin v
c con ngi.
3. Thi cc cht giu dinh dng cho cc vi sinh vt bin.
4. S nung nng v nc thi nhim nhit t cc nh my in.
b) nhim c th lu di hoc ngn hn. cho thun tin, ngi ta chia
nhng nh hng nhim thnh nh hng di hn v ngn hn. nhim thng
xuyn bao gm s a vo cht liu c hoc nhng nhn t chu tc ng ca con
ngi khc thng lin tc v mc thp r rt, gy nn s suy thoi mi trng. u
vo ca cc cht dinh dng bt ngun t cht thi (nc cng) l mt v d in hnh.
u vo ngn hn (short term) c nh hng r rng nhng chng c th tiu tan vi
thi gian ngn. Trn du l mt v d v u vo ngn hn. u tin, du gy nn
nhng nh hng nghim trng i vi vng sinh vt bin, nhng nhng nh hng
dn dn c ci thin ...
c) S nhim c th bt ngun t nhng ngun x hoc t loi nhng im
a l. Trong mt vi trng hp, nhim n t ngun x nh mt ng ng cng
rnh hoc ca cng nc thi nh my. Nhng trng hp nh vy tng i n gin
trong nhn bit v qun l v cc c quan iu tit c th tm thy ngun x v gim st
quy m, phm vi khng gian ca cc nh hng . Ngc li, nh hng ngun phn
tn c th l do bt k mt v tr no . Dng chy theo ma l mt v d in hnh v
ngun phn tn.
d) Nhng nh hng ca cc hot ng con ngi kh c th nh gi mt
cch khch quan, nhng nhng s thay i trong tnh a dng ni chung, s c mt v
cc iu kin sinh l hc ca cc loi bioassay v sc chu ng .... c th c dng
cho vic kim sot. Tnh a dng cc loi c th c tnh v mt s lng cc loi v
bi thnh phn phong ph. Tnh a dng tng nu s lng cc loi tng hoc nu s
giu c v loi c phn b u hn. V d, mi trng sng vi s a dng thp ch
gm c mt s loi vi u th tri hn hn bi mt loi.

Hnh 10-3. S suy gim a dng cc loi ng vt y c kch thc ln
188

(a) theo s tng nng hydrocarbon trong trm tch gn khu vc khai thc du (b)
theo s tng nng ca cht thi dioxit titan trong vng bin Nauy. H
S
l php o s
a dng tng s loi v s cn bng tng i ca phong ph cc loi.

Cc cht c hi
Khi cc sinh vt bin b nhim cht c, cht c c th khng tng v nng
hoc c th tng theo thi gian trong c th sinh vt. Mt s cht c tng v nng
trong cc sinh vt khi nhng c t c vn chuyn thng qua mng li thc n.
1)- Nhng kim loi c: kim loi thng tch lu nhng c t v c nh hng mnh
m khi con ngi tiu th chng. Hng nghn nm, cc loi kim loi khc nhau
c thi vo mi trng bin bi cc hot ng ca con ngi. Khai thc m l ngun
kim loi ch yu v m ca sng l ngun nhim chnh. Chng ch yu l thu
ngn, camidi, ch.
2-) Thuc tr su thng c dng git cht loi cc su b, cn trng sng trn mt
t nhng chng b ra tri vo trong nc bin v thng xuyn gy c i vi s
sng bin. Thuc tr su bao gm nhiu loi hp cht khc nhau c s dng
git nhng cn trng, su b c hi i vi ma mng. V s ph hoi ca su bnh
thng phm vi rng, hu ht cc thuc tr su c ng dng mt khi lng ln
(gn 1 t kg trong mt nm M) v rt nhiu loi khc nhau (c khong hng nghn
loi). Mc d mt s loi qu thc c hiu lc song hu ht cc loi su bnh u c
kh nng khng thuc, loi thuc tr su c hi nht l DDT, dieldrin, chlordane.
3)- PCBs, nhng loi ny xut pht t cc hot ng cng nghip v c chng
minh a ra vn c t chnh trong mi trng ca sng. PCBs l loi hp cht
c s dng rng ri nh cc loi du nhn khc nhau ca cc my mc cng nghip.
Khp ni trn th gii, PCBs c thi xung cc b bin v c tm thy nh mt
cht gy nhim ca nhng ng vt khng xng sng, c v loi ng vt c v
bin. PCBs gy nn ung th biu m chut v v th n xem nh l nguy him i vi
loi ngi. Nhng vt cht ny c bit l mt vn v c tnh rt cao ca chng v
trng thi n nh ho hc. Mc d cc vi khun bin c kh nng phn hu PCBs song
qu trnh ny xy ra rt chm.
4)- nhim du: Loi ny c c nhng nh hng ngn hn v nh hng ko di ln
cc qun x v cc loi c th. Vo 50 mi nm trc, nhim du tr thnh mt
vn chnh vng b bin. Khoan du, vn chuyn v t chy, tt c dn ti vic
gia tng du vo mi trng bin. Sau y l nhng ngun nhim du chnh: (1) R
r t nhng thit b chuyn ch trn bin v cng, (2) r r t nhng l khoan du, (3)
r r t nhng tu ch du b v v xung, (4) ra tri du t nhng vng dn c ra
bin. Mt s nh hng phc tp ca s c trn du c nghin cu bi cc nh
khoa hc ti Vin nghin cu nhit i Smithsonian sau s c trn du trn bin
Caribe Panama vo nm 1986. (Hnh 10-4). Rt may, vnh ni xy ra - Bahia Las
Minas c iu tra trong nhiu nm trc khi trn du trn b. Ban u, trn du

ph hu xp x 7 phn trm din tch rng ngp mn trn di t ven b. Tnh trng suy
tn ca cc cy rng ngp mn khin chng b nh bt ra khi v pha bin, gy ph
hu sinh vt y. Du cng ph hu cc bi c bin. Trm tch khng lu na din ra
v vn chuyn v pha bin, tng c v qu trnh lng ng trm tch v lm gim
s sng st ca cc rng san h gn . Du thm vo cc ht trm tch rng ngp mn
nhng ma ma n sm v bo, dng chy nc ngt lm du tr nn n mn v vn
chuyn ra ngoi khi, lm ph hu nhiu hn ti c bin v rng san h. Thng thng
h sinh thi rng ngp mn gi cc ht trm tch gip lc nc v gip ch cho cc
ngm san h.

Vng gn b

Trm tch

Du

Rng ngp mn
C bin
Trm tch
Xi l

Rn san h
Xi l do
bo
Vng ngoi khi
Hnh 10-4: Hu qu do trn du nm 1986 dc b bin Bahia Las Minas Panama.

Ngun dinh dng v s ph dng

1)- Cc hot ng nng nghip v nc thi lun mang cht dinh dng cng nh cc
sinh vt gy bnh vo ngun nc. Cc hot ng nng nghip v s pht thi cc cht
thi gy nn s ph hu to ln i vi s sng vng bin v lm nhim bnh c v tm
cua. Tc ng ch yu l gia tng cc cht dinh dng v lm gim trc tip cht lng
nc. Cc vi sinh vt khng mong mun b phng thch vo mi trng bin t cht
thi cng rnh v cht thi ng vt. Nhng mm bnh nh virut vim gan v nhng vi
khun Salmonella thng tp trung bi sc vt n nh con trai v c th l nguyn
nhn ca nhiu loi bnh. S bng n bnh l v cc bnh khc c th tr nn ph bin
nhng vng con ngi thu lm tm cua gn cc ca cng rnh v nhng vng b
nhim nng vi cc b cha nc nhim khun (gy thi). Cc c quan mi trng a
phng thng tnh s lng cc vi khun coliform trong nc bin. Nhng vi khun
ny gn kt vo nhau trong rut ca con ngi v chng c coi l c lin quan n
nhng mm bnh khc. Nhng vn gn y ng Bc nc M vi vic vt b
cc ph thi thuc y hc vo i dng lm tng kh nng nhim cc loi bnh cc
k hi (c virut).
2)- Ngun dinh dng, cc hot ng ca con ngi dn n s gia tng cc ngun
dinh dng ho tan trong nc bin. Hin tng ph dng l s b sung cc cht dinh
190

dng vo nc, dn n lm tng s sinh trng ca cc phiu sinh vt ph du ... v
hot ng ca vi khun. Khi nhng ao nc ngt cha y trm tch, hin tng ph
dng xut hin mt cch t nhin do dng chy v qu trnh mc nt thc vt thnh
nhng vt th nc tng i nh. bin, hin tng ph dng do mt s ngun dinh
dng lin quan n cc hot ng ca con ngi. Sau y l cc ngun chnh:
Ngun im nh cc ca ng x l nc thi
Ngun im nh cc h thng ng nc thi
Thuc tr su s dng mnh m cc vng t nng nghip v dng chy b
sung thm mt khi lng ln ion nitorat v amoniac vo nc mt v nc ngm
Xc thi ng vt l ngun ng gp nit ng k trong dng chy c bit l
nhng vng nng nghip nng thn.
Hnh 10-5 hin th s ng gp s b sung nit n lu vc vnh Chesapeak.
Kh quyn c th l ngun dinh dng chnh b sung vo vnh b bin. Gn
y, ngun Nit mi c tha nhn l ngun ng gp ng k vo vng bin ca
cc nc cng nghip: t chy nhin liu ho thch l ngun pht thi oxitnit tim
tng chnh. S pht thi nhng kh ny cui cng c quay tr li tri t khi nitrat
ho tan trong ma. Cc vt cht tr thnh mt phn ca ma axit vi thnh phn sulfur
c th lm gim gi tr pH mt s cc con knh v ca sng ti c i vi c.
Nitrat lng ng bi ma c th gy nn mi lo lng bi v chng c th kch thch s
sn xut s cp gy nn hin tng ph dng.

Hnh 10-5: S phn b tng i ca nit Vnh Chesapeak

nhim nhit
Cc trm pht in cn nc to nhit v l nguyn nhn dn n s dit
vong ca cc loi hi sn. Trm nng lng ht nhn v trm pht in thng cn mt
khi lng ln nc vn chuyn nhit t h thng pht in. Hai cch thc tiu hao
191

nhit c s dng. Trong cch thc th nht, nc lnh c ly t vnh hoc sng, i
qua thm thc vt, nc nng sau c lun chuyn thng qua thp lm lnh ln.
Sau cc thp ta nhit thnh cc kh quyn ph ln, nc lnh li quay tr v sng
hoc n c th c phc hi trong mt h thng khp kn gia thit b nng lng v
thp lm lnh. Cch thc chnh th hai i hi nc chuyn tip t (ly i) t vnh hoc
ca sng, i qua thm thc vt vo cc nh my pht in v nc quay tr li mi
trng tng i nng. Thng thng, nc c dn quay tr v h nhn to sau
mi chy ra vng ca sng hoc ven bin gn vi nhit cao hn nhit mi
trng xung quanh khong 10-15
0
C.
Pht thi nhit cng c th nh hng n nng sut cy trng, nhim nhit
hn ch sinh trng sinh vt ph du v thay i c tnh qun x thc vt ti nhng ca
sng m v vng ln cn.
10.3.2 S khai thc qu mc
Nhng tc ng ca con ngi v s khai thc qu mc cc loi c v mi
trng sng ca c c cp vi nhng vn sau:
Vn vi khi nim nng sut bn vng ti a v s suy gim nhng mng
li c nui
Chin dch qun l cng nghip c nhm t c nng sut bn vng cc i.
Nhng xut gn y v khi nim nng sut bn vng ti a c th khng gip ch
i vi vic qun l ngnh cng nghip c. Khi nim nng sut bn vng ti a bao
hm quan h c bit gia sinh trng qun th v ln qun th.
1)- nh bt c qu mc gy nn s thay i chnh trong cu trc dinh dng ca ct
nc. T do vo bin v s tr cp cc hm i nh bt c ca chnh ph trn khp th
gii dn n s sp rng ln cc cc ngnh cng nghip c khc nhau.
2)- Gim ngun cung cp c th do nhng s thay i ngu nhin hoc t s bin i
mi trng, nh bt c qu mc cng c th c coi l mt nguyn nhn.

Hnh 10-6: S gia tng s lng c trng (ng t) v s suy gim c xac-in Thi
Bnh Dng (ng lin) ngoi khi vng bin California.

192

3)- Khai thc c voi qu mc: khai thc c voi bt u ven bin v sau pht trin
ra ngoi bin khi. nh bt c voi cng l mt trong nhng n lc ca con ngi. Mt
s k thut tin tin c p dng trong nh bt c voi bin Nam cc.
Ngoi cc vn nu trn, cc n c c th b nh hng do con ngi lm suy
gim cht lng nc hoc mi trng sng ca c, hoc do git trc tip m khng
phi l nh bt.
10.4 Bo tn bin v h sinh thi bin
Vic bo v v s bo tn b bin v h sinh thi bin l mt vn ln, khng
th cp chi tit y v vy phn ny ch a ra mt s chin lc v bin php
chung lin quan n vn ny.
10.4.1. i vi vng b bin
Nh cp phn 3.9.1, gim cc tc ng ca con ngi ti h sinh thi
b bin, chng ta phi c nhng chin lc trong s pht trin v qun l cho ton b
vng, t thng lu sng cho n vng bin
1. Pht trin cng nghip v cht lng mi trng. Cng nghip vng ven bin nn
c t nhng v tr c nhng tc ng ln mi trng sng l nh nht v khng
lm nhim n nc. Vic quy hoch cho s pht trin ngnh cng nghip nn bao
gm nhng vng cng nghip, cng v phng tin tu thuyn. Bo co tc ng
mi trng phi kt hp vi cc ch tiu s dng bn vng.
2. S chuyn i rng ngp mn. S phn vng nn c nguyn tc ch o r
rng chn la nhng vng cho bo tn, bo v v pht trin. nh gi ti nguyn
bao gm gi thay th (ph tn thay th) cn phi tnh n khi lp quyt nh. Gio
dc cng ng cn thit nng cao hnh nh, gi tr t ngp nc. Mi trng
sng rng ngp mn nn tnh n trong vic qun l cc k hoch s dng bn vng
hoc bo v
3. Nui tm v ngh nui trng thu sn khc. Vic s dng mi trng sng ca
rng ngp mn cho thu sn cn c xem xt li. Phn vng s dng t, cht
lng nc cho nui trng thu sn v tc ng mi trng ca thu sn cn c
cp trong k hoch pht trin quc gia v a phng.
4. Bo v rn san h: Vic tun theo lut nh bnc vng san h l cn thit. Chng
trnh gio dc v s tham gia ca cng ng nhm cng c sc chu ng a
phng v la chn cc hot ng ph hu cn c lm t u.
5. Cht lng nc. Vic a ra cht tiu chun cht lng l cn thit. S hp thnh
mt h thng qun l lu vc v sng nn c thc hin, nu c th vi kim sot
nhim vng cao. Ly mu v kim tra cn c tiu chun ho. Nhng ngnh
cng nghip v cc ngnh vi phm tiu chun cn phi c kim sot (iu chnh).
6. Xi l b bin v lng ng trm tch. Cc bin php kim sot cn phi c thc
hin nhng vng c nhng h sinh thi qu gi rn san h, c bin, rng ngp

mn, ca sng v b bin b nh hng bi xi l v bi lng. Nhng d n xy
dng gn bin nn c nh gi tc ng mi trng. Nn chm dt hoc c nhng
iu chnh s khai thc m v no vt dc b bin hoc trn nhng rn ngm san h.
7. Pht trin du lch. M rng hng dn qun l mi trng v lu lng cht thi,
ngm san h v cc h sinh thi khc... nhng vng chung gn du lch. Cn hn ch
s thay i x hi.
8. Nng lc v s sp xp cc t chc. Cc c quan c quyn hn qun l vng bin
v cc c nhn c o to cn phn tch cc vn qun l v pht trin cc k
hoch. Mi cp chnh quyn cn tham gia vo ICZM. S tham gia ca cc cng
ng v cc t chc NGOs s ci thin th ch.
Mi quan tm qua cc phng tin truyn thng. Sinh thi b bin v s bo tn
cn c ng h bng cc phng tin truyn thng i chng v a vo
chng trnh gio dc nhm nng cao s nhn thc ca cng ng.
Cng c php ch. Cc lut l hin hnh v qun l b bin cn c xem xt
k cng v ci thin cho ph hp vi thc t v c th c thi hnh, cc lut l
khng c tc dng cn phi c loi b.
10.4.2. Bo v sinh thi bin
1)- Bo v bin khi nhim do cc hot ng ca con ngi nh cc hot ng
pht trin t ni a ( lu lng nc thi), vng bin (du lch, cc ngnh cng
nghip...) v trn bin (khoan du, hng hi...).
2)- Khu bo tn bin l vng a l c thit k bo v cc cng ng ch
yu v cung cp ni sinh sn cho c vi hy vng chng sdn phn tn ra nhng vung
rng hn. V s tc ng ca con ngi v p lc nh bt c tng khp i dng, c
bit l dc b bin, cc qun x bin ngy cng b xo trn cng vi s mt n nh
ca cu trc mi trng sng v cc loi. Chc chn l khng th chm dt cc tc
ng mi ni nhng cn a ra mt s iu chnh thch hp, t cc vng ct yu
c th sm c bo v.
Vic thnh lp cc vng bo tn bin c th c iu chnh t hai quan im:
Bo v nhng vng thit yu duy tr v thm ch m rng qun th cc loi c.
Bo v cc mi trng sng thuc cu trc khc nhau nh ngm san h hoc
cc qun x khc.
3)-Cc khu bo tn bin cn phi da vo khi nim "metapopulation" (siu
cng ng). Mt s qun th a phng c th tr nn qun th ngun m cc c th
t nhng qun th a phng chuyn n cc qun th a phng khc bi s lan
rng cc u trng (trng nc).
4)- Mi trng sng c cu trc thng xuyn b e do bi s s dng ca con
ngi v c bit l i vi du lch. Do vy cn phi c hng lot chin lc bo v.
Mi trng sng c cu trc thng ph thuc vo s duy tr lp nn n nh.

5)- Vng khng c php nh bt bin cn c bo v tt nht. Khi nim
khng nh bt c th p dng cho nhng vng ct yu a li mt c hi nh nht
nhm duy tr ngnh nh bt hi sn bng vic cho php cc qun th hon thnh c
mt chu k sng. (Hnh 10-7).

Hnh 10-7: Thit k mt vng khng c nh bt mt khu nh bt c theo l
thuyt. Vng dnh cho trng v c con v mt vng cho c trng thnh l
vng khng c nh bt, cho php mt qun th nh nht c th hon thnh c
chu k sng. Cc vng khc cho php c nh bt, mc d lng nh bt phi
trong mt gii hn cho php.
10.5 cc h sinh thi v mi trng ven bin in hnh
vit nam
10.5.1 Bin Vit Nam
Bin ng ca Vit Nam c din tch 3.447.000 km
2
, l bin ln th ba ca th
gii v l mt phn ca Thi Bnh Dng. Bin ng c hai vnh ln l Vnh Bc B
(150.000 km
2
) v vnh Thi Lan (462.000 km
2
). B bin Vit Nam t Mi Ngc
(Qung Ninh) n H Tin (Kin Giang) di 3260 km. Dc theo b bin min Trung
(t H Tnh n C Mau) c ti hng ngn hn o ln nh.
Phn thm lc a ca bin ng ( su t 0 - 200 m) c din tch 2.640.000
km
2
chim khong 76% tng din tch b mt bin ng. Phn sn lc a pha ty
bin ng c cu to bi nhiu bc vi cao khc nhau, sn lc a pha ng ca
195

bin ng c b rng khng qu 74 km vi nhiu bc v cc h su m st qun o
Philipin. Phn y i dng ca Bin ng kh bng phng v c su trung bnh
3290 m.
Nhit trung bnh ca bin ng khong 19-27
O
C vo ma ng v 29
O
C vo
ma h. mui trung bnh ca khi nc mt ca bin ng l 32%o. Thy triu
bin ng kh phc tp v c 4 loi thy triu phn b cc ni khc nhau (nht
triu, nht triu khng u, bn nht triu v bn nht triu khng u).
10.5.2 Ti nguyn bin Vit Nam
Ti nguyn bin Vit Nam gm 2 nhm chnh, l: (i)- ti nguyn sinh vt; v
(ii)- ti nguyn khong sn. Ngoi ra bin nc ta cn c ti nguyn mui v cc iu
kin rt tt cho pht trin du lch.
1. Ti nguyn sinh vt:
Nhm ny gm thc vt v ng vt, trong bao gm ng thc vt cp thp v
cp cao.
V thc vt cp thp, bin ng c khong 653 loi thc vt y (ch yu l
rong, to), thc vt ni bin nc ta mi ch bit khong 103 loi to vng
vnh Bc B v khong 300 loi cc vng bin pha Nam. Theo tnh ton ca
cc nh nghin cu th trong vng thm lc a nc ta c sn lng thc vt ni
khong 1000 - 1400 triu tn/nm.
Thc vt cp cao bin Vit Nam ch yu l rng ngp mn, chng phn b
ba khu vc chnh: (i)- Bc B (t Mng Ci n Thanh Ho) vi din tch
khong 80.000 ha; (ii)- Trung B (Thanh Ho n Vng Tu), rng phn b ri
rc vi quy m khng ln, ch yu cc m ph ven bin, cc loi cy chnh l
c, mm, s, vt,...; (iii)- khu vc Nam B (t Vng Tu n Kin Giang),
rng ngp mn Nam B c nh gi l ng th hai trn th gii v s lng
ging loi (sau Malaixia), din tch khong 329.000 ha.
ng vt cp thp, theo iu tra ng vt ni ca bin ng c 710 loi trong
110 b ca 13 ngnh ng vt khng xng sng, trong c gi tr nht l
mc. Theo cc nghin cu th khi lng ng vt ni bin ng trong tng
nc 0-100 m l hn 3 triu tn v c xu th gim theo chiu su. ng vt y
bin ng cng rt phong ph bao gm san h, cc loi c, trai s, tm cua,
hi sm,... Trong ring san h c 303 loi san h , 200 loi san h sng
v san h mm, chng phn b b ty vnh Bc B, cc o v qun o nh
Hong Sa, Trng Sa v ven bin min Trung.
Cc ng vt bc cao bin ng ch yu l c, b st, th c v v chim bin.
Ring c l loi ng vt cp cao c gi trnh kinh t ln nht, Vit Nam c 4
vng c chnh l: vnh Bc B (c 961 loi c); bin Trung B c 50 loi; vng
ng Nam B c ti 2000 loi vi khong 100 loi l i tng khai thc chnh;
vng Ty Nam B c khong 1000 loi. Tr lng c cc loi ca vng bin Vit

Nam t ti khong hn 2,769 triu tn. B st bin ng ch yu l ra v
rn bin (c 3 loi ra v 10 loi rn bin). Vng bin Vit Nam c khong 200
loi chim thuc cc nhm hi u, b nng, r, mng bin, yn,... Th c v
thng gp l c voi, c heo nhng khng nhiu.
2. Ti nguyn khong sn:
Vng bin thuc ch quyn Vit Nam hin pht hin 4 loi khong sn, l
titan, t him, ct thy tinh v du kh. Trong Titan c tr lng khong 22 triu
tn, t him c tr lng khong hn 300 ngn tn, ct thy tinh th c tr lng rt
ln cc vng ven bin min Trung. Ring du kh l mt ti nguyn chin lc,
chng ta pht hin trong vng bin nc ta c 8 b trm tch cha du kh, trong
mt s b ang c khai thc nh Cu Long, Nam Cn Sn.
Ngoi ra ven bin nc ta cn ti nguyn mui c th khai thc hu nh khp
ni dc theo chiu di 3260 km b bin, nhng tp trung ch yu min Trung ni c
mn cao v nhit cao trong thi gian di trong nm. Cnh quan p vi cc vng
vnh, cc hn o, bi ct,... dc theo b bin cng l mt iu kin rt tt cho pht
trin du lch ca nc ta (vnh H Long, Sm Sn, Nha Trang, Mi N, Vng Tu, H
Tin,...). Nhng khu t ngp nc ven bin cn l nhng khu vc c h sinh thi c
gi tr cao trong nghin cu v bo v mi trng.
10.5.3 Nhng h sinh thi v khu vc mi trng ven bin in hnh
Dc theo chiu di b bin nc ta c hng chc khu vc mi trng v h sinh
thi ng vai tr quan trng cho pht trin kinh t, x hi t nui trng nh bt hi
sn, giao thng, nng nghip, du lch,...n bo v b bin chng xi mn, m bo an
ninh quc phng,... Trong mt s khu vc in hnh nht nh sau.
1. Khu vc vnh H Long
Vnh H Long l di sn thin nhin th gii c UNESCO xp hng, H
Long vng bin ng bc Vit Nam, thuc tnh Qung Ninh vi din tch khu vc l
5937 km
2
. Vnh H Long c din tch rng ngp mn khong 3600 ha (1995), c
nhiu h sinh thi vi tnh a dng sinh hc cao nh h sinh thi ven bin (littoral
ecoystem), h sinh thi o (island ecoystem), h sinh thi gn b bin (sublittoral
ecoystem), h sinh thi c bin (seagrass ecoystem).
Tnh a dng sinh hc ca cc h sinh thi vnh H Long l kh cao, tng cng
c ti 810 loi ng, thc vt khc nhau, trong nhiu nht l thc vt ph du
(phytoplankton) 177 loi, t nht l c bin 1 loi, c c 109 loi, san h c 143 loi, v
cy ngp mn 15 loi,... Khu vc ny hin ang c quan tm bo v, song do qu
trnh pht trin kinh t x hi mnh m nn cc h sinh thi v mi trng ang b tc
ng (khai thc than, pht trin th, du lch, khai thc hi sn,...)


2. Khu vc ca sng Hng
Khu vc ny bao gm cc ca sng chnh Thi Bnh, Tr L v Ba Lt thuc cc
tnh Thi Bnh v Nam nh. Din tch khong gn 100.000 ha, trong din tch bi
t l 25.000 ha. c bit trong khu vc c din tch rng cy ngp mn t nhin v
trng khong 3000 - 4000 ha.
H thc vt khu vc ca sng Hng kh phong ph. Thc vt thy sinh c cc
loi nh lau, sy, ci, rong to (to l ngun thc n ng vt nc). Thc vt trn cn
chu thy triu ch yu l cy ngp mn (c 6 loi) vi mt cao 5000 - 6000 cy/
ha, trn ct c mung bin v sam bin. H ng vt gm c ng vt ph du (c ti
185 loi) ng vai tr rt quan trng trong vng ca sng ven bin, l ngun thc n
cho cc loi ng vt khc. ng vt y ch yu gm cc loi hai mnh v v mt s
loi tm. C c ti 156 loi, trong c khong 40 loi c gi tr kinh t. Ngoi ra khu
vc ny cn c nhiu loi chim sng vng rng ngp mn v chim di c theo ma.
Sinh thi v mi trng khu vc ny v ang c quan tm bo v (c khu
bo tn Xun Thy), nhng p lc pht trin kinh t x hi vn l mi e do thng
xuyn (gim ph sa, thuc tr su, khai thc qu mc,...)
3. Khu vc ph Tam Giang- Cu Hai v ca sng Hng
Ph Tam Giang- Cu Hai v ca sng Hng l in hnh ca cc ca sng
min Trung, vi chiu di 60 km ch rng nht 3 km, hp nht 0,5 km v din tch
khong 22.000 ha, su 1,5 - 2,0 m.
H thc vt nc c 9 loi cy vi kh nng thch nghi v pht trin rt tt, c
ti 86 loi to c pht hin (c nc ngt v nc mn). ng vt nc c ti hn
100 loi c thuc 11 lp v 54 h. Ngoi ra trong ph cn c mt s loi tm c gi tr
kinh t cao nh Penacus indicus, Penacus Monodon. Mi trng v sinh thi vng Ph
Tam Giang - Cu Hai v ca sng Hng ph thuc rt ln vo s khai thc ngun
nc sng Hng, s hot ng ca cc ca sng nh Thun An, T Hin,...
4- Khu vc ca sng Tin (ng bng sng Cu Long)
Ca sng Tin thuc tnh Bn Tre (mt trong 9 ca ca sng Cu Long), khu
vc ny c mt din tch vng ca sng ln ti 25.000 ha. H sinh thi khu vc ny rt
phong ph, c bit trong c ti hn 16.000 ha t rng ngp mn, y l mt iu
kin rt tt cho cc h sinh thi v bo v t bi t.
Cc loi cy loi ph bin ca rng ngp mn l tra, mm, s, vt,...v cc loi
thc vt ph du c 278 loi vi mt ln ti 100.000- 3.000.000 c th trong 1 m
3

nc. ng vt ph du c 36 loi, trong nhiu nht l ng vt gip xc (c 26
loi), ng vt y c 24 h, ngoi ra cn c nhiu loi tm nc l sinh sng trong
vng. Ring c khu vc ca sng Tin v ven bin c ti 43 loi bao gm cc loi c
nc l, nc mn, v c c nc ngt vng ca sng.

Mi trng khu vc ca sng Tin rt thch hp cho cc h sinh thi pht trin,
c bit l rng ngp mn nh ngun ph sa phong ph. y l vng c tim nng ln
cho pht trin kinh t. Tuy nhin vng ny lun chu s chi phi ca dng chy sng
Cu Long, thy triu v cc hot ng kinh t ca con ngi, c bit trong nhiu nm
qua rng ngp mn b suy gim do khai thc qu mc v do pht trin nui trng thy
sn.
10.5.4 Nhng vn thc t v mi trng v sinh thi ven bin Vit Nam
1. Nhng hot ng pht trin nh hng ti vng ca sng, ven bin:
(1)- Khai hoang ln bin, ph rng ngp mn ly ci v lm m nui trng thy
sn, canh tc nng nghip.
(2)- Khai thc ti nguyn vng thng lu, c bit s dng ngun nc cc sng
(h cha iu tit, p dng, trm bm ti v cp nc).
(3)- Pht trin nng nghip, nui trng thy sn nhng vng gn bin (m ph),
xy dng cc bn cng vn chuyn nguyn liu, pht trin du lch cc bi tm,
thnh ph ven bin.
(4)- Cng nghip ho v th ho cc vng ven sng, b bin v pht trin cng
nghip khai thc du kh, vn chuyn du kh bng ng thy,...
Cc hot ng pht trin kinh t c nh hng ng k nht n vng ca sng
ven bin l nhng hot ng nh hng n lng nc, cht lng nc n v nhng
hot ng lm suy gim din tch v cht lng rng ngp mn.
2. Nhng vn thc t v mi trng v sinh thi vng ven bin Vit Nam
(1)- Suy gim din tch, cht lng thm thc vt v rng ngp mn do cht ph
rng ngp mn ly cht t v lm ni nui trng hi sn, quai ln bin ngt
ho vng ven bin. V d, din tch rng ngp mn ca Bn Tre gim khong 40%
trong nhng nm 80 ca th k 20, nay ang hi phc nhng chm; hay ti ng bng
sng Hng, my chc nm qua do quai ln bin mt hng ngn hecta rng ngp
mn. Mt rng ngp mn dn n xi l b bin (Cn Gi, thnh ph H Ch Minh),
gia tng nhim nc, gim nng sut sinh hc,...
(2)- Gim ph sa v ngun dinh dng t cc sng do nhiu nguyn nhn, trong
c cc h cha, p dng ln nh h Ho Bnh, Tr An, p Thch Nham. Cc v d,
p Ho Bnh lm mt nhiu bi ca cc n c kinh t vng nc l v bin
nng trong khu vc ca sng Hng (theo V Trung Tng). Hay lng nc x t h
Tr An trong ma kit c kh nng lm gim mn ln nht khong 20-30% v thi
gian duy tr mn cho php no cng ngn i nn nh hng n sinh thi vng
ca sng (Cn Gi). p dng Thch Nham ly ht nc trong ma cn, lm nhim
vng ca sng, mt cc loi thy sn c gi tr,...
(3)- nhim vng ca sng, ven bin nc ta ang l vn ng quan tm do
cc nguyn nhn nh cht lng cc h sinh thi gim, ngun nhim t nng nghip,

nc thi sinh hot v cng nghip ven bin, nc thi t nui trng thy sn, cc hot
ng kinh t khc ven bin,...
(4)- Mt s vng b xi l hoc bi lp nghim trng vi xu th ngy cng tng
nh hng n n nh dn c, pht trin kinh t, n giao thng thy,...V d xi l,
bi lng nghim trng khu vc b bin Thun An- Ho Dun- T Hin (Tha Thin
Hu), Mi N (Bnh Thun), ca sng Ci Phan Rang,...
(5)- S ng vt y vng ven bin nc ta ang c xu th gim, c bit l
nhng loi c gi tr kinh t cao nh tm hm, vm xanh,...nguyn nhn l do khai
thc qu mc. Mt s loi c cng gim v nh bt bng nhng phng thc hu dit
nh cht n.
3- Nhng kin ngh bo v sinh thi vng ven bin:
bo v v to iu kin pht trin bn vng mi trng v cc h sinh thi
vng ca sng ven bin nc ta cn thc hin:
1- Nghin cu ton din cc quy lut din bin ca cc iu kin t nhin v mi
trng trc khi thc hin cc d n pht trin trong vng (dng chy sng ngi,
dng triu, cht lng nc,...).
2- Khi phc v trng rng ngp mn ven bin, c bit l nhng vng d b xi l v
nhng vng trc y tng c rng ngp mn.
3- Quy hoch chuyn i c cu kinh t nng nghip vng ven bin hp l, xy dng
cc chng trnh quc gia v bo v mi trng v sinh thi vng ca sng ven
bin.
4- Nghim cm nh bt hi sn bng cc phng php hu dit, nng cao nhn thc
v mi trng v sinh thi bin cho cng ng ven bin, ng thi tng cng cc
phng tin hin i cho nh bt hi sn xa b ni c sn lng c cao.
10.6 Cu hi tho lun

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