the plankton: drifters of the open ocean · figure 15.7 zooplankton sample (enlarged)....

15The Plankton: Drifters ofthe Open Ocean

n.1 1M Ki"",oIP1-kt...

""rl...•...-~

Lh1_f(....a-J f~

IS .l II---.IS.) O_lKM.....s.--,.oIot."'-"'n.• ~."' P1....b_

[ .... Yr...... ;" ... 0..-..

IS.S "-'i..l~,.... M...... T.....RedT _O......T Saoo-Cipo P";~._-

K" T"••.......S.XP'IwIR~

-__'Ii"'__""_

I

I ~ • .....lkn .wUt:nin~. ina<d ihl< in ita owift I!I< sm.,.....pi..... 01 rho_1><w;in .., "'....,""", ll>e;. ; 01ao.""""",nJ 1""J'O'1;""" Th" tpri"' .... bdooKo .. lint to ....

Ji."..... """ to all ,"" on... ",0:,,..,,,.. fMD' lit. ufdo::plonkton.ln ,"" fJe.t« "'.....1)'"r ,hoi, gn-lh .beyroo<:<""•.-ofO<nn ";th • r"'"lI bIonkct of!beiJ clls. ~iI< all..",iI< 01 , •...-. =I Of hro....n or gner>, II>< """""...n.. '" on If>< mIor 01 ;nfl.ni,";,N1 gnoi... ofpiwnrn. <t>I'I"';-' in -" "' plw. <dk

n.. piul.. "'.. undiopu'ed ......y in 'h< .... lOronly •....... ,i_.AJ_ .. ona ,h<it-.. bunt 01,,,uj' ipli<o,,,,,, i•....td>od by onc=o< in t ho: .....u anin>ili of ,I><......k_ It ;. ","",u,. ...... ofm."''M'''''and ,Ixr-- ~ <h<impand ri>< ";ngcd oiI. Hunm___ 01, bltl< bNsu oIlhc pbnb_ n'1 "''''''&'> do<....... tioediow ,hot ohwo&n. pIoft" and riocmsdoo:s UIli,.1""1.., ......c--..

.....eo.-&- n. s...k...J U.

1'"l'lIlCtll'" '''' ' l> [o r," IM~ " Ill,II ,. wh'ch pl' ." ..I,' ,10 <'1 ,,·,

a"d '-'""1,,,,. ,, wi'k V;' ''~fr "r C>r~ ' "l1 ''" ' ' i , ,, hr< I ,, , ~ !'i ,1I."d c,,,h,, 'The ' r ,,,' i,,li,,·,1 , )I'~ "" i ",,, In,,,,01 I,,''' '" ." ,I.,!i<" R"nu ," ," ,," OC, II" I\owl"' ,,, ~ I ,,'

G",,,p' 01 orp ''' ' ''' ' hclollg , ,,~ '" ,I ,~ 1,1" -- ' ''I'l:LIl I.' "",ncl"do , h ~ ,li"I ""' " '!i,,,,f1agt"il,,' ,', , " ,,,I "" , . " .lilh" p h" ,..; , il i",n,,~d l ,"e,. cryp",,, ",,, ,,,h ,-t ". ,,,. ,,,, ,""",, ; ' e.", alg"". ",>""""""", .'1' '" hl"" _."" ,.·,, .,1,· ..,n 1''''''' ''' h., Ie" " ",,"',"", . II 1"" )('", I".·,l l'h" ,.'I,,,,,,,, ,,"""plt .. <how" i" hp",: '-'. 1 In tl", rd l ,, ~; ,,~ ,1",·"".<>",h. trill'''''''''''' I~;,,"d ",' ,he " ,," ..m, .",,1 ,I", ,1i," ,Il",· .·1I"'e•. a< Ihey arc ,I>< tn,,,, """ ...1" ,,, , ,,~ I ' '' ',, ' "nl~ " " ' '' '

" >t,,,b.... " f tt>< ",",,,,,, It'y"'~I,tn k".,.

Di.""", "'C ,",~I.- ,d"'~ ~I.",,, 1" ,,,,,1 '" .,r~.•' "Icold . ",,,,-i.II'·rich .. "'~•. I'he)' ;or" ""'0.."",,,, , •.,Iled ."" 1,.·,,,I~""" ",:<:a"", ,loci. ch;oc.K'"ri" i,· y,,1~ "., . '" " "" I~ ~rl~·",

r~","a", hi", ,,~, ""'i, chl""'phyll S.",'" .1;",.. ,,, I""rad ia l '~ B1 "" ' <y: 'Io<r Jr" """'d. , I,.,po;,1 hh' 1~1Il,,, ,,"

.,01 n,e ~a:I<,1 ..... , .->< ";a"~n" O.h<o-< ....... hil .," ·,·" I " ",.

"'tlfr , a,c· clo"!,-;,,,· , ;,.... ;I(~ ullod I"' n n ~ l . <li"I''' '''c .... ,;c ,IWo".. n"", Ioc>IC' ,1>.", I"''''''''~ .-1'."1\"", .I" ,d . ~.'1"'''''"'''' d..to"" .,~ "lI ,'" r",,,... on ,I>,.. "'.,I1,ow "·,,n.," ,~

, " "" hed '" fl,",' ;,,~ 01';":" . "h,Io; ,,~~.I( "';"'" II' ".~ ",' . '(ruly p l'n"onic St",,,, ",M",""" di;,hM' I",,,, "''''1-·'' '' '..'". " ." . ,11""" ,n h~",,,, I~ 1 "",1" ,

A,oo,><I ,he "",,;,,,-" " f e.och di""~,, ".' I, mt,d. ,, ~

c-.:II w.1I1 .r pc<;""'."r ~ y hl·• • • ' '''-''' y,I, ."c . i '"",-~;""J. dw;,h , i l;<'• . Th . f , . " . lo " h" '" '; ~.. I "" " '1' .'0<'" "n,1

"'ii"" 15, /",,, "opI~ , t ' '" ' [>,••,,", ," . " ...." ,,: "".....".'....." ,. , ' ~,..'"••, "",_.<c. II• •,.,~."", •

15.1

Phy top lankfonTh. phy,opl, n .'"" .~ ", . ,,,Iy ""'<'<'lIul.. ( O< " "8Ie celle<1 )pl, nl< kn" wn ...Ig. c. E. ch ph'·'"pl," k,,,,, ce ll .. . .._1<>1'01'" ;" (", ""I f recd ill~) hy 'h. p"" :<.,, <If I"""" ' yll' '''''i,(rtfer ,,"ck to chaplc, H ). E;\\;h ""II i••n ",d<JIO"<lc"' i,KI, ·.id". I. " KI ~.~n i" ,h. '1"", io. ' " whi,·h the «II. all""l, '0gothor in r;ram~lI l" (1""1' , h. in,) or ", h~,- ,,"'"" ""'l<.,t><rc IS n" d,vlS'O" of I" I~" ' 1"'1 " the « II" TM.." .. ".. Iy01>< large p l.llk' '''' '• • 18° , Ih woc,1S"j'~u""", th.. i.lound noa l in~ in th" arca (If tho N"'th Atl.ntic kllOw" ""Ih. S.,I'''-'"'' Se• . SUI'8i1"SU'" ' 0l', ,,duco. vCI'eu, ivoly hy

"' kl>o"~h many pl. nklOO line . I,mil, d , t>ilj,y I" "'(lVe I" ·

.... d all<! '''.r from !he ...~ .." I...'. , ,I",y m.ke no ru l'J'O' . ·fuI m<>lion .!t"''''' the <>;:U<l', <urTen" ,,~j .,. <."iod f"'>1llpi..,. h> pIaee ' 1L'f'""'Ied i~ the ,....w. let . S<>m< pl"~lk)1I " '~

qu;le la,ge; je llyr..h m.y be lhe ' iLe " I • lOl l\< w. dllub....,I'rIll 15 m (:'>0 Il ) " I 'en".d ... II " , ,... phy1''l''''nk'oo .,><1.....y roupIankloo..." g<nefally 100 ' 111,11 rn, ..." un. ..",edV,"IOIl and mu<l be oboe<ved """",, • n'M;ro><:opc,

Sac",,,••,od -":" 11 ph l'~lon~", " cell> .rc c. lIed"II'.pl.a"kl"" ; ' ''''y .., I 1_ 0.005 mm in d,.",e"'t .,ldcan ooly he e-,I1 ",,'ed ",ing ' p«,.1 (,I'e" ng IOc hniq...,"SIiSM y 1..,..- phyroplonk""', called n."nnJlI.n~t"". ha••

• >lU " lIge betwe<:n O.flO') and 0.07 mm. 7.ooplraIlUo " ,"dphytoplanUon belween 0 .07 .~d I mm .... ca lled mi·ct"plall~'''" '" lIel pl." kl" o. bee ", t""y ate ...... lIy cop-ruTtd in low Del. m"de of """)' rm. , "yloo.

TIle m,cm«,,!,ic ptIylopl,"k'on "'~ Ihe ~g,....... "rIhe ... ... )" .. " " 1....1 wi'h",,' gnu _ he"'" could "'"' upl"-'" 'he i" ...". ,m.1I ""","n" . " nd hi nl>. ,.......tv<! ..

food 10' lhe 1"8er. ""',...' a' in& cami . ",,,,, . • _ ,","boulphy 'opl. II ~'''''' could nOl ,uppuol the """,,1_'00 iIO(l 'he01he1 IMII. r . nim.I•. A. (he Btlli"" bloIus",.1~.

ph« Si, AI" ' ''' fl.,-dy I.., '"'d . ..All I1e!h .. g'''', H

T bt ......,.d f/,,~/"'m "'>N'fr- Ih<c............... pl.nktoo,."aning I. "",nJ". and IIx""'~I.""n, ,",,, I"-dl...

au llu wand",,,, and J' 'f1m ifllx "" , TINy ",ill i """" 'm,, hmited ,. Ibn' "'.b,';". ,"""ins ""'h ,I;, ,.n~nJ,

TI" d,v",,, IJ ./ pianl:. lOn" "K"m'''', " wXm ,I I"'1 ,/ " not1"'"i/;/, tQ dmY" 1111¢ft lmr 'tft-ftrm' h<>,. {nJt...J, ...~" .

lali'll< "tt, ,,,,,I<,pI""", 0"'/ g'.~!,' ofam",,,I, ond".,,/, Nwh«tt ,,,,,,,,,fin' J,,,"won, R«""" " " Ii>< plant lift I"'" " .hkf. M" ,/)< ,~~ i <n"K)' left' '" Ih. btl,,,allift j'" "lIrht an,,,,,,,,,"'" b-lI;n by"",,,aninx tb, ..-,an ', fl""t '''l pl. nr 'oft, 11><,n""'ry 1'''''''u,., of' hafl" 14. " ,,,l lh.. "" On 10 tI"'ri b< lIN.m",tlill;";' drift ""'h Iht p'"nt!"'n' ton, I!.ra~inx "I"'" il 'Iff'!'1""'4tb .'!Mr.

:JI!C The Kinds of'Plankron

..

f'r,,/S,1n.. """_ _ ,""""'_....-'" ..."","S ' t _ : y r......l .. . ~O:lo . ....yof_ '.I T... _ r-'_i'~. (b) n.. ¥'',__ ~,... _ '·"lA"._ ..... (~) .......; . i lK .."'_ " of 4"",." ,' ...... " ~ I~)T", /"l<"ft'I ,,,,,,,,,, -_ ........... "~....... of "~ ....... ~".""""i"

~I

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, "\•• •

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, ,• ••

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,

•

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.,

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f'ir ..U.l~'~."

4•• ..- " ,.. _ .._.._~ ,., or_lfl C_...._,_or_ w""' .....-

"

Fip 'Y15.JC-"" ............. • •_ Il,............. -"' .." . '" m ""',,"

o,s .....

Fip,., H. ls.a.;.- ..,,""'... .,._ ,_,,'"

<ll;li<o..ly m.. l <d "," h p<>rn III'" ronB«;1,t>i"" 1<> ," ""'lS,<Io en" u"nma>' ( 1'3 ' 1' ,0,n.. ' ''''0 hal... of . u nlric J i. ,om', f' o.role fi' lOp'''''h k~ • 1',1 ' 1><>. , ",d ",!>en 'h. cell ha' ~m",," ." rfic i". ,lylarge , 11'1< « II inside d,,,irk,. lhe ''',0 halue, of ,PIe pill"'"sep.,"te, " "" ... inn<r holf 10 eac h p,IIb.->< i' fo'nted , . nd,...0 He"" d. ughle' d,.,om, ore prodlLced. Hy 'h" p""",,".0"" of Ill< ,leW coil' will remain ,lie ...O\e ""' "' ,he p.'len l,whib< ,h. ", he, i. ,I ", . y' . m.Ue, . "' i l> r.r~e, pjlll>o. h.lfis ronned by ,he >l1\I11e, 1:>00100' half of " '" p"'e'" pinho• .

lb. pnx::e.;s .. $IIo"'n .. f,P'"" 15.5. W"'n attll ....,"'" aso", Ie""l of "booM Z5'1o of~",,~,ouI 1"'1aII" . ia; il ""I'>d..idLn~ ..,d , iM a ",.oa l cycle. In ,~" cyde ;' pt~.

" ""ken .0"" , whicb ,n,,,,,,.,,,. In sill'. f ."-",,, a ......f'"S IUle, . "d bell'''' h' d...de "gat" . D"'om, di..de "e,y,"p,dly, e"",y ''''due 10 I""enty-f,>u, hours o"de ' G""d,ll" n"of pleo', !'u l ' o" ha lol ""~ nutrients, When thi, rapid dov i.i""ItIC",a, e, ' he l"'I 'o la' i,.. <Q thou the w.re, be<:omc< di",,,I.need by '~ I'r."'""" "f mil"n", and mill]on, of rell., ;t i.coiled 0 hlno", .

f,K"" lJ.5Tho di' i' ''''' 01. ~".. ,~" , n< d.."", . " " ".'(,d"."..- ,j ,_m~ TO. ''''' ", I." Qr " ~..,,,,,,,t,,, " II .."...~ . I I~ "II <c> '"""d" .... ""' . ... ~ I,.... " Ir" '''''''d lor ,....,",,_ .o..~, ",II ,.., '''" '', '''' .. '.. p"'''' tI.""" "",.11",

f 'K-.. lf. 6~ __ G_"'--,...,G ,,,,,,,,,,, 1"'><1"", ~d ,.... 1'1.. ",,,,~ .. . _""'"....,,"', "'''''0' '''"1,""<1." 11,', ,C"",,,,,I,,,,,,..' od~ -.,. _'" ,........ p"al" 1I1h."'''''''''t:

I'ru..u.... made wi'h " liea 0'" """" de""" 'hwl ..a·w. .... "", ,lit d,.,,,,,, muSl .,.y alloal in ,II< ,u"li, .."f....,W"'" '" """,i.e. 11>< m..mal ""II "",... ial h... "- den ·"'y and...,eaocs i" ",",oyancy by ,he producti"" of oil 0"s ln..,e prOO''''I; f'''' Ih. , leed Oft ••' l:C qd...Ii';", or d, ·~''''n. may "",e a di. lonaly oily '''''''. 1lIe d,.."",· , , ,,,,,II,"u ab o help' 'I I" Slay . no.l, becao.. ""an p. nocln ,h... re <M, I~' <yore ' "'POnded " n~ ¥cry , lo",ly, A ,,,,.11. ao~ In alorgc. "{'hcncal particle ha, . lorgc ... rrae<: . "'. ,n oompari·""" 1o i" , olum" or m.,,,, and thi. l o,~c "",f",e "C' '0volu m~ ,.tio help' keep the cell aflo"- Son-.. d,.,on.. f">S'""" . pin," , wing" 0' " ,her r mjeC UofUl th. t 111<0=" the", urr. co 'rea still more, A IMge , " tf.cc an:a . 1", p, ,,,,j,",,the d'. lOms wilh increased expo' ur" t" , ,,nlii hl ' nd I"wOle, conloioiog me ooccs," 'y g.,e, and o" lrienl> f 'lf ph"lo.ynlhe, i, . "" growth.

DiOioms a,., nm' l i" 'p<lrl"MI a, the fin l 10' 01"I' f(~>d

(lloo"o<Iion . Th,, '" Iha' an: nOI <nn, umed by hc'hi'of<'e.elll"ally d,e and ' ink 1o !he "" e<ln 11<",,- 10 , haIlM'" a,,,a,of Ihe oce an" Ih. <~II, reo<h ,h.... n oor w,, 11 , ,, ,,,.

<>Ii""" n",,'~' "ill I"",kcd i"'ide, [)eros'!' ", . <1e long as"m 'hi< man"" , h' ''e f""""d fM'1ro ~"m. 0' oil. Dialom 1'1'0'·luI<> ,ink , n; IO '~ grea'er deplh, of 'I>e ocean, blJild ap"liccoo' scd'men" "ntk. are"" of . oooo\;l n' d,al"''' popIll..Ii",,, (Ref.. "","k '0 ,e.. ""' loon on bio""""", <ed, m. nlS in<,,"pIeI' 2.) Somclimc. geologic; procc...,. loll lhe... "I",..,odI 1it:d,n ~••bovc ,he sea. ..-hen: ' hoy . '" mlflCd ao d O.~'..n' n"". ..h",h .. ".... ,n i"d,,,, . i. 1 f,I"."""' y",co". ,n ,he f,lIe, ing of ...in< ., "cll ., ,"'immin ,pool" .1Ida. an ahrasj"", in lOOt!l....le _ . ilvc.-lJOI"h

The dinof1.g~Il.'e, din •• from ,he d"."..... in ... . .~r.1 ,e'l>ecl" Dinoflagellalc, "",all) ha.e hvo na~.lIa. or"'hiplok. appe"dage.. ,10.. heal ",i lh,o 1l":MWCS ,n lilt "" II.... 11 One 1I,,,,we enc;TCI"" , h~ cc ll like . hell. ~,M1 ,heOIht"T lio, al ' ighl "'lI"" to il. The healing ot '''''''' nall" lI.n,"le. Ihe cells """ ,k . nd co""" lhem '" ",in hke 1"9" "Ilt e ~ mo"o Ihrougll 11t< w" l~f They .1,,, ,."d '" mi~"'''

'~rl i.. lly i" ,e' p<lM'" 10 '"Mlight. 00' ,hi• •bi(ilf I" m"""" li"' ''oJ. and thoy . re sli11 . r Ihe morey of lhe ...""••,,~

com nt' . Repre,enraL i, e dillof l,gc llate' . ... , ho"''' in r 'l·u... 15.6

Figure 15.7Zooplankton sample (enlarged).

Dinoflagellates are red to green in color and can existat lower light levels than diatoms, because they can bothphotosynthesize like a plant and ingest organic material likeall animal. They have both autotrophic and heterotrophicabilities. Heterotrophic organisms feed 011 other organismsor on organic substances. Their external walls do not contain silica, hut may be armored with plates of cellulose, giving them the appearance of spinning armored helmets.Other dinoflagellates have a smooth, flexible outer surfaceshowing no such plate structures. Some dinoflagellates arecalled fire algae, because they glow with bioluminescenceat night. (Refer back to chapter 13 for a discussion of thisphenomenon.)

Dinoflagellates are found over most of the oceans butdo not contribute to the bottom sediments, because both thecell walls and the soft parts decay completely. Althoughthey do make up a substantial portion or the phytoplankton,dinoflagellates are not as important as the diatoms as a primary ocean food source. The cells reproduce by a divisionprocess similar to that found in diatoms, but without the reduction in size. They can, under favorable conditions, multiply even more rapidly than diatoms to form blooms.

Coccolithophores and silicoflagellates are relatives ofthe diatoms. The coccolithophores are single-celled plantswith coccoliths, or outer calcareous plates, that are deposited

as sediment when the cells die (see fig. 2.19b). These coccoliths are soluble under the conditions of low temperature andhigh pressure found in the deep oceans; therefore, calcareouscoccolith deposits are limited to shallow regions of less than4000 m (13,200 ft). Coccolithophcres are also limited by thesurface-water temperatures to the warm tropic regions, andso coccolith deposits are to be found in shallow areas at the.lower latitudes. Like the dinoflagellates, they possess twoflagella, and like both diatoms and dinoflagellates, they mayreproduce by simple division, although some species do havea form of sexual reproduction. Silicoflagellates are small autotrophic cells with flagella and an internal hollow skeletonof silica. They are not believed to occur in large numbers.

ZooplanktonThe animal members of the plankton, or the zooplankton, areeither grazers on phytoplankton (herbivores), feeders onother members of the zooplankton (carnivores), or feeders onboth plants and animals (omnivores). A general zooplanktonsample is shown in figure 15.7. Many of the zooplanktonhave some ability to swim and can even dart rapidly overshort distances in pursuit of prey or to escape from predators.They may move vertically in the water column, but they arestill at the mercy of the waves and currents and so are considered planktonic (or drifting) organisms.

Tbe Kinds ofPlankton 373

.:>

iIIII

f •,f ipn U ,fIr-_.........._ .....__.. (L..~.~.( <• •,~,~,'... ."-" ....

R."......IIU'l i..... 01 _ Iy ~.c'Y ..,....1ph, '"

f".",d 1M zoopa-~",. '"'" I,j" II"","" of """" "Itt.! ~ "" 1M lO<Jl'I ,"" _ •.ned MId uncl yuI . ,.-,_ f 1 "' I ..or ld ..tic ,.., reprud"" " 011,...,..., h"l1 ...:1 M. sr- _"",,,, n...., ."m. l, y~~ to , ,_ ta'C"',.... . year In "'. rm "'., ......... food ouppI... at< __ POd1o''''I''''''u,,, -=-Ir.·... I' le prn<~ At "'~h 1.",_. ",llorc l~' • ...,., ' 0'p/lyt<>pI..~_ . ,,,,,,'11" bn.f, ,he ,,,,,,,I_~ 1On ml 11"0due. onl,. l ill,l. ......ou".. in . yur , The yorociou."J'P"UIa and ...,,<1 110*,11 "'''' of ,... ,"'""or",,, 100'pUn kLon lnpOIISible rm lobo•• '; ". ,,"ui......o b. I«y'. led~ , ptlyl<oplank'on

"l.uopllnkl"" ,,,,,, ,1\ ,,"'d"" <>f h,~h pupull'iooJ dc:n.." tIc,\W<:n I,..,,,,,, ,..... In: m",,11 I. ....... '1' ''''I'''laI...J ,11w hi,lI """,,101""" rotehe>. Ml.oc' predal<>n. Ind 1M rnun:'J"'~ popul.,~_ 1><, ........ ,tic dense, 1*. ..... Pf<'I"O"" 1M>lock , •• re""". p<Wo"'" fO<d 'hon'. T"rt>u1en<c and eold,"d,'1"". ,nd,.Kl...I. f",... 1M ..",...1, """",..... """' 0lhe ,n'<"c..," , .p• ....,• •",u. C<>II'. ' 8<"<0 ' ''nC' <1""undo,,,,, be'\O'ttn ""0< ly J'C" " lflCCn" '" ..><>pl""kl<WlI"'P"'- hi<;h . ,lrael precb<on.

1'l..,U ac;.....m<J . ... .. Ihoc: olmo;"y bouodo,lOo ..-lby 1IIt la ren~, of lI>e , .., fa<.'e ",alen . _ ,hr _ o f

liltbl " ilb deplb and .... doy ·. i&b' < ~d. play ,,,'.....1role>. Some zooplonklDrl ml~" . to..... ,d ' hr "a ,(_

••d . n'I '" ...,. ..,,,rn 10 ""I" b udI riay• •i'.... UI an ."."""

'0 ......nlaln ,""i,"tb' I..... u ,n relf'OI"" '0 !be "",• • ,_

o f .be rt fuod .C>DU'•• . Th, . da,ly m' I '"l '" '' ",ay be atm"d, ••~m ( 16:10 I'll OJ I... t"'~ 10 ... (33 III~"",,,I.,,,,... 01"'PO""" In . 'bin bond .,tend",&

b"nton,.lly .1""," py<:noo::lirle u .1 a pt t k m:d kltbl in"",-."y ,or f d ..SOUl.. Ic\<d . .. UlI"'bIc "I pao".It~ ..fIC<:trnlt......nd 1 fronl dtpcb ""',....... Tht zooplri"'" llye '.. _n "n a Ntby""'tri. rec""',n,l' a falte bal",,,, or Id<ep ' <I n. t'''S II~", . tb. DSL. (Ref.. '0 . h pt. , 4,)E<bo- sound'''l .."di.. "'" ....... 10 record the ........ miIt''lion of Ill ,. lay... 01 plankl"" """~,... ud to tnClI_

' u'" ,''' _',col _ bun"",,'aI e.1ct>l of ,Ill: ' . ye"A"""" the nlD&l <omo_ oJesprcod n>upI.onIl -

' on lypco .....nd...KI< ""' .... ,,,,a n <T (IIIrim"hkc' . "" 01.1' n opcpod tul'!>. .., i id. (fi,. IS &l. n.....c ..,_mil'" ..... bo., ....lIy rt.-. and """"' more than holtlhe rr _ y ","', lit dl,ly Copcp<><!> Her ,h"" COl·plooooo'odo.: eup/Iauo.iioh ....,... more "" 11 .nd bV<: l""lUtho" ,be c"l'CP"'h , Tht tupllou' ''df . """'au... of t""" ,i,•...... u ' ....... 01 ,"" .....Ilcr rooplankton .10"1 "lib ,""pIlylop1ank_ th.. malo ~ ,"" .... 'k of therr (oud. Ilolb re·prndu<:c __h ",on: lIo....y lh.m the ",.,001S. doublrn, ' ....i'1"'P"10,,0,," only 'b"", ", f""r lIrt101 0 1"0.. l1lo:y ....y "'akeup mon; than 6f"'- "'!he zoopl/Ink tOll In any of the .. ukl·,",U n' . nd "",v<: .. a f"",1 "'. for .... an f"h . In IheAIt ' IC Ifld Ant..... "'. "'" ."p ""," .re ,I>.: kri ll. (X;Curri"g,n ouch 4""'''''' ,hat they I'w.,dc U•• "" i" lood for lhe

Fit'"' 15.9no. ,,""",., ""( E__.....,...,."

bal«n ..ha l~•. lIaJee" (0' ...h. I. OOn.) ..h. I•• h... nnt..,lh; ,n" . od t h<~ h.... n. ,Iok< w.,,,., "f h, l..n , ,,••poeoood from ,I>< ,oot, "I ,hoir mouth, . An., the ""'h,,lo.gulp tho ... 'er arod pian. '''',. , It. ~ ."J..I tlo....te, t)uou~k

,be balttn . 1.3Vl!lg tfle tiny krill bohinrl. Wh. lr . or Ihi. I ~poe

inch.to Ibo hll.... " l!.hi. humpb;1<;k. ",i. anJ fmhock ..hal..T I>c Ant."'ti. 1<.,,1 1. E"I,lwu .,iu ''''1'' '''''' (fil . I~ .u ).

are 1"""'''' m e,,,,,,,,o,,, ~" . "I'l«" Eo'""... . ,~ un. 11>10_"""" have vaned from ~ mill1n" 'n ~ b,lI",n ,...trie 'nn •.and aro probabl y in ..ce.. of 'XlO m,ll",n met" . I" n).B«ause of tbe", large biOlrw• ••lIn,.l« , ,Ill: .,, 11 in 11>0Soulbo,n 0.. .... . '" co",t<le",d. pule,,' i.Uy ..1""",1e inler""t""'al 1..1Il:1'~ Thoy .... h............. cn<nmore,all ~ by I,<h·,ng n..t, from ,h. f",mer USSR. Jap.an. "'''"'•. Poland.. n<I Chile. In 19~1 ...1 1982 Japan anJ lJlc fom..... Su'<",tUni"". u. ingl,up: .lem " ...10.. "' ,Ih ,"". !Ill m (2M rt)

..ode and c:aldl,n, 8-U Iono nl krill ,n • ""lie haul. ,,""."".led ...,g.(l{ll.,,,,,,io; ' ''''' ,n 19111 _ ~29~.(l n.....'" ."".,n 1982

Marl<1onl ,ho: loll lor """'... COfI'Un'pl'on h., ooth<>::n ...., """,.oJol. F_ . "n arc: ,t_ n oowheIl dried tho: " n Ot becomes ..,"'" _ -..e h•• Un_

pk»onL The ....11 ""' bu. f......... '0 <o'''a'n •.at ",mel ~

hogh ...........1. of n "",ode .... m"" be rml"".d I", humonCOI"<OO,mpl_ In lIlo: f...",... s.... Un ,.... the ...<h .... Ictl

....,n1y 10 Io.-n>od pnul''1' , Japonc.., uw It.." ." ll

.. 1_ on ....... r f In . d."""" ""' .. ,, 'he ..poe' '''''<IiIk k>nf: <howa In ttw:. fiohinl " 1'10<;, thoboll ll<fcr,...... ",podly. lIlo:y m.... k I""""' d b< n........ Iomll,nl lIlo: da,ly hMW.t 8« of the ,lLrr",ul-lies lIlo: ~.ill <;on "".opo:l bo:1_ 1 ~ 1 ..... I "'K~ to Ie..lhan 2OO~)()(I _t"" loou . n<I '"~ lhcn to.. .~.'"ll'" ....Ihln 400.000 .... tril; 'on' . Thr I~ <I 'eh ..... 3~ 1.()110

,_,,'" 'on•••I'er 1100 bo ,,!' of ' hr s..vOet 11",,," 'h.cM<h IJtopp<d 10 2It9JXl'1 " '" 1<.... dun"llhI: An''''<1'''1991_ 'n_

AIl ,h. na""". p"" u lIy h. ... ' " "11 k,,11 ,n ,""Southl:m (),e.ln a", n..",he" " ' Ih. ('"nvrn,,"" f<>f ' hrC"""""' ali"" of A", ."·,,,, Mar"," !.iv,,,, R......n.a. and .nn.•• all'''"'' ,,'" ' " "....,......,,, III.. ""')(I' cr. 1"'-_...,e"lu"""" of Ih. knit r....",. ..,~""'" In to "' ,,"', ....., 1hI1

...-h," tho krill "" I •• ,lable 'n ....1t" nllml><"" ' hey"" lbe1000:1 of mon~ .....ales...... they I""" • "".ic link m the londweb lor tho .., .01•• pal", i"• •001 00>er h, ..... >quid. ...:I fl ..,

of ,hi: A",."" Oe "'~""'; ..... tbc boo in "';, chlpor,Arro .....onn•• or <h"" " ,!= U lh. (ICC r'll. 13 ~)••••

•bond.n ' in ""• • " "" '.'" 1m ... lhe .."f""" 10 the ' ..... 1dcpIb•. lllr.., mocmocopw; (2-3 em) (l 'n )• ......ty "''''9''' 'onl. • " ,acious c.m,~OI'O' recti on otho: , ....mbe'" " I ,hezoopll nkton. s.c.e,"1.'PC'<"'" of ."_ _ no .'" I"""" .ntho ""~ . arid in '<"'11: QOC• • p"" iwl. 'l'""' iel. .. lound <wyin . <co,." W'O<' ,n .." . n... .."""",,"" ........... ..~..n. nd w..... m...... ... """'pIrI. Ih., th< .pecies earl b< . oed,,, ,drnl,fy ttor "rigtn of ,ho: wa'.r ..mplc tn wINc h ,I iI

f" u..... In tho NOI1h Atl.nt... tho: a""",worm. Silt"'" u ,,,''',, ,,,h. hlt' only 1hc t>oorth s.. ....e< ....... and S rlrK<"'Ji, f""",1only 'nocranH: ... '<n-

.·"r~ ,,,i ,, ih·r. n• •nd . .... ioobor ilJ'" _ .."""""~.

"nlll • .o;;el led, a...""t>oltk. prOlozoanS; lbe, ........."'.. ,nl illU'. 1) .10 . f o.am,n,lua".....<h as lbe cOmmonr;ta"i/t""'''. are .......Kd tn • """'-ponrnen uka.-.co-;,,,n,.'" ohc1l, ..hil. "'" ndIoIanano ..., nd<d b)'

• "loea Itfol. 0' ""' II .The ,aJ,,,Iari"'!em ""..,oIyorolptu' .... and """"~ wi,h deliQoe 'PI""'. ("'"'''~. in,hi: """ atl<"" a """~""'l)' """"""" ttIIemal pr............ _an ..",mol l.Y'" of 1"",,,,,I...m .~.. (f.h" I_I,m. ny wol" . h lel. 1 .d. men.., radI'''' oul ff"'" lhe <ell.R.~"I.oiI". foed 01\ d••t""" and .mall pl'tllIWJo .....hI ,"llle. poeuJ,>pnJi• . Ilolh f."..mon,I... .....nd .aJ' a l." .....'" ,.,..... ,n lIlo: .....""" "'~'''''' of ,ho "".-. AI....""",h .lho......11. """ tr'" """um"I. ,e on ,he ""un fI ,,,,,,.•nn·I" hu""" kI lhe . edllnem•. Calc."""" f",,,,,,, n,fe' a" le.1Sare fouOO ,n . 11.110..• "'. ""di....n" ....h,1e Il>o . ilicwu..oJ""..' "'.... ...luch ani I" rbe d'....,I.,,,'" ";,i",,of the 11:,. 1""'<10<"" ""• •1 i"""t., tleplh. , commonlybe"'''' 400f1 m (l J .200 Il) (",. eh' ple, 2. l ig•. 2.19 • .,d2.2l)) l'inh nnl<l. 1_ fiB. I~ . 10) .'" tiny 1"'>\<"1>0'" withm"" i,,~ 1I."I ,k" . trllelu' ... . ar <lli•. The'" ,or".,,,,"" .." flO" <11I.d hell • .,,,,..1, , ",I 0,.. f" ",,d ' " eo....1 wat....

""" in ,he "f'On " ,:"",1'le." I"..I. (f ,,, . I~ 11) are l1Iul l",'" ,h.y I'" ",laIod

10 It.. ,,,. iI, .not . IUIl'. Thr y moy 01 may "'~ ~ ••o . , mall.,.k:am'u, , hell, d<po"d,"" 01\ ,he '[1«;". but .11 h... af,,,,, 'hal" ,"ndir.r<1 i,,' o ~ tran'p. ",,,l . nd gracefully "ndo·latin, wln~. TI'e" h.,,-d, eak ..e"", ""n.in. ennu,!>u,. to' he bun"", nd im.n l, in ,h. II" ... tropic ' ."'001 Sum.I~rtopod. orc ho,h,.", .. and """" .", <am,~"""

Tun'pHe n'. dehcatr. and tu m, ne OCe" l, thecI<nuphorn, no . noM ,.11 ,., (fig. ". 12). nOl' in lbe ... ·face ",.1. ", . Some have 'rail.." "'""",1.-.; all d e JlI"POllcd. lo",ly by cii!>' . "w , ,,( belt ing <iii• . 'The U. •oundf....... .... famili. ly callod ....,,~ "' lnu'"by <0llU"". tho ,,",unf,,1. tropical. "''''''''. lLJlI<:ncol Ven...'" rd l< may , row 1<> 30 em el2 in) '" m",e ,n lenglh. "f1'OUP nf V...... · t'rdla d"I' ill& "' the ",If_ and <"", lItn,tho: ...nll~ ","k the" hea" nl <illl is. """"!.l<ulat .."h'hom II"" da: k of. >!lip. All cte""""""".", ,"nivor-..,I.....,"ll .... CIlber """l'l...kton

•

"'r-H.l'_-......._\.'10_ .. ... _" ". "W.'.'__•__~_ ._

llot h..out<. ....."..,. lrompuenl __ <Allie u>o

piW..Io>II. II "'laird 10 1M noore ""'lII<cd .en.bo_ ..,_...b { ~h 1wo<~~J~ II> 'oodpoIol,~< 1M

u l I"""" sal 1_ ",. 1~ .121 "'" pel..,,," , it...,. """~ cyllfld" ,, '" n.......-aH; 'ho, an: ,,""" ,y f_ in__~ ",..1Cn:d~ ......y _ t. ,... ufloU-<-BOlh , " ..... l'tIatoc nlftOUln• • ~_p. ",1-Iyl l t e and " 1"'.-. • .., IIOl 10 be: .""r...... w"n .l" ll yfi'"(h,_ I$.Ill. T Jellyf.... """'" Irom allOlhef ..... ,,""'btedlrouP'" .,mah, ,hot O~ItIH ...t . or C·Ilk/ori• . Some )Clly''''' . Suc h U In" common A",.I;o ond ,n" <01" " ,,1C"._ WI'" II. !roiling .. ,n~,nl ""'laCl... ..,....t ,I><'" e,,·Ii.., 1,_ .. dnhen. 00 &lOCh ... C"",OIt<''''''' .• "",. 11jrllyf.... til n.. Atl....oc , Px ,foc a<e......nd A<<J_ N ,

founJ in ...." y "''''PC''''''' "' "",,,,hot.. of ,... pl..~ ,

1M fo<only . pu<1_ '" ."".. 1,_ 'hoy ~""'u.lly "",,,Ie..... cha>v 10. booom <1... 111"1. • lIac""'l f,.", """I., w •....... ...._. A""'M' 1'''''' 0' un"...1,.lIyli,h .'" ,'''

"'P'. 15 11n.. ..._ .. •

Fip" H.11Tht _ j!fl;" (...."'......) ~......... _ _ . I'fo........._ <>Ilod ._..........~_, ~_

. .. I.. " ia l f.. r m. , in(;llJ\l,n, ,b. p_~,....... lIlan·o f ·war.f'~y.><Ij,<l . and llIe ......11 by.,"".w.I'Id....nlot. Vrklla Bolh

.., roll....,"",.. <If ,rId,vldual but "'P"Cia~itod ... ,....10. """'"

of ....hidl ....~ Iho tuk <If 'alh.....,l food. ~''',. orpn>Iect '''1 ,ht roIon) .."h .. in" n_cello. wtI"k """'" 'orm• Roo..

A ll <If "'" zoopbnl "'" d.""....""" '0 ,hi> ""''''. wil!lthe u <qlbOI\ <Iforrta,n ,.1I,r"...'l"'"o1 ,heir mli~ Ii."' ....p lo"k1oo and ... ca lled bolopb "kln" (see ri,. 1~,14)

Ho__. an .mpon. n' pori''''' of fhe zoopl. nk'oo """ "I,onI, pan 01 M...... pl.nl'on. ltoe.e a~ "",,,,he.. of the_ropI ll T1>t me,opl_h>n ,,,,,1..01< ,he eu.', 1.,.••1

ond ,... ,'" "'1'" 01 rna") ,.-_",,,,,," "".11 ' I"'"~ mo" of,he" h..,. a• • i'be, f", • . ...."n.... " t, "",h e, li, h) III i>ot l<l lll

d....e1Ie.. (..,,"!l ... ..robo . nd . "u fi. h). 1'0" f. .... ..... . k' . thel' '' ;0<1 (".. '''''"~ IMm,) "f"Y""", 01""•. 1.., IlIl< It-' . c...b, .wornu., ",0,1 . . .. . ,-fj,h•• ,,<1 Ill ;' '' y " ,be, '''ll. ni."".re a panof ,he ""~)I""ln>n. 11", ..urrclll. " noy ,he.. I.moe ,,,..,....("""'""''' ",", ,< ,t",y r" ,<1 ... . , 'n """I. and food ......n.:e<

Jln 'h .. way. re-popul.,,,,,, of . ,... or, .... I"" h • , pa; i•• m,\,

......, doodOIl' <1<""" .and o"'''' lm'Idin" n ,h. h,>tI'" "re. i.mluad Se ,mar. I"<><IU<_ I.m.. 1l\ _ ,,,. .. . " .. ,,,,,,,00,,.and .... I ropl""kl' ''' . ,. on "np"'''""h. ,.J "" ,,,:(: fm0It- meonbo" of ,he w op l''''t,., . utl ,,' her o,,,m , I•. The""Tefl' . ",m. l. ,n. y prod uce ", illion . or .I' n. nu' <>n' ,, m,lI " "mbe" of m, l"" , ,,d fomol. , ,,« e. fi", l~, I ~ . nd I ~ . 16) 1'....1, ...."",i" ho r"""d .",1 ,10"",,10«1 lheoo I.",. ,...e oa<h •,lO , ' h,,,l.inR the~ ...., d,""o. _I1><1 . ..... ' YPC 01 ani",o!We kccop WI"" of ,_ """.... lOII. y• .., fom ",. , .pie . '0 II... lruchoplll>R' •• "'. ", .....,.,'. ,.... ~Iil ..of <e• •noil•. ,loc """" I. ..... of en"". ....1 ,loc ~, 1.·v.. oftw..... l..

Olhe, ,,,e,,,ben of , loc .....""'Ionk ' .... ;,,<lwdc r,W.egg•. I e• •nd Jt... ",,," f.. h. n.. y....nJ ri. h Iced "".......... I "". uOI ,1 <hey IJI- r... _ Jh 10_ ,...- _

~.••

~.

".

\

• •••

•••

\

r~1J1I__..... ...c-..._..... . _~.. I' • _4 _

___ ........._101 ...'__·'· ....__..

.-11I ..........

•~...

....I...... ' U'______ l .'

__., - .. .. 1 ""'• •- ..

,.

l ,oc_ . ........,"'", , ' , -rnj

~ '" ..........,,,"' ....'"File", H .I . ""'.An.., ......... r",,,,,......""""' ... ,,,. """".....

""

~

f'Op ... ,Ut.-_01 ..s ..__...._ .. _ ......... POO'" ,,,. .............." " I flor ..... of . ....... _ l '~l'" .... ", . ........k·I ..__• • _ .. . ......w. ,

A Krill-Rased Ecosystem

AI..-, .11 Ih. 101. " I An"relic. "r><I,he .um"'nd,nt Sou 'hern (.J,'..n Jepends"" the 'Ie.>. TIl< ',,00 web of the SOI"I\e'~

o.:un " I,. ..d uO t ~c phy top lankto n,1n.",ly d..lnm" ,oar h" n. " Ihe ene'gyollh. " " .. F,, "n'. te> of I'~mary prOOuct;on ""y ; 1n " "l1me,. ;n ice·free are• • ,tiLl' ..or'gt Ii." be,w""" 20 on<! 100 g ofc tT bo n V., . ~ ". r. meter pe, )."(r C/m2!y. .. ). A ,non~ ,h . h.rbivorou,"""pl,nk, ,,,, ,,,... ,peei,.•• Eup!>.u, i. ",.pert... or' kn ll, prnbobly MOUU"t'> t" !>allthe tOl,1 ""'Pl."t"" h,om.., . K,ill .'"the k.y <><" "i" ,,, o f ,he S<:>uohem 0«: ..""""1"'01.' ..." , ,~.. 'ncllOdes ,he . , .comn'.""I) " r or,....." .. and lhe etIviron·"" ,,, I .."h wh",h ,. 'n..... o.. Allh"" gh~Tlll , .. c",· ..",poIa ,n d i",, "",;"". I"'i,<on<.n".II"n i' not un,h"n, • • nd thc~1"lnl o"n, """. , io" ""e~ .. in 'h<W<tldcll .'leo. hel_" 0" "od 6O"W, S,,,,,,,

' w' nn' ...."" been "" ;II ''' IUI "more th. n2 ""II"", \<.0",

In , unm"" , '0. K" I'''d mel" beck,the ,..itn."y P' oO 'K-ri"n in"" . ,.. , • • nd the~, L1 1 mO'. Im," " I, h. , o,;oe< 10 groze {I",

phylopl.nkLon 0• • 1' I ,,~. A"",. In wint."pr;m" y pr<>d""tiun ;, p" ," " . 11 1 1l00 .from rho .0 '1'..... wAl.,~ d.. '0 ..".,"«1Ii~h t , i, '• .p."~ cO~r• • nd ",cR'oed tu ,·b"l.ne~ ; "Ihi , ,i"",. 11.. k" U .~.,

'0 de<o:.. ", 'f'lIU <It<per ....... omJ feed ""lb. ph110p l.n~l "n "''' '' u, . "The .0n".1p,o"ho tlun o f k"n bi um , n Ih.Suu'be, o Oc.... ha. bu "mal.~ "1'iO million---lJOlI m,lho.. ''''''

.""'uld , .. ao i_,.n, PO" "f II",AIOl'''''ic food .. . b, The,~ ....~ he """"dun ' ''''''''~ 'PC"ies. \01,,,, del"'",I,n~ nnkll ll .. Ib.i, rn<:><' impa " ' ''' fwd. The'0,. 1," " ",1 co ,,,u," p' i,, . of "l ,, ;d bywbol... b ird" a"" , • • 1, i' cak" lat. d "' bou' 35 m,lIi"" ,,,., S""'. ' I"' '' '' ' ofAnta",hc (i,h <L' y 111 ,h. S",,,ht,rn lkean) ear.wund, Ie<dm~ "" ' ,i ll: ",I",,,"",.;1.,nug" ,e lnt" A,,"ret;e W"e'~ ....·b " ""me, to rm.! "" . ,, 11 1, 1" , heon e,,,m"N

'hot oil ".I,,"'i, (';,h ",nb;ned m.y C"" ,

,,,nle ,, ''''''' h", I ' ~J milh"" " '" of hal..ell y••r.

n ....,. I. .. ,pee ie, of lim" i"A.ta L'Cli"• • bOI por" l.. i",,, . ,., u,ually, ...y "'1<" R"d< of A,• •"·.i,,, f~ 1'fl'I"ipall ~ On c,o>l.c•• 10I.io11 h i ll .....eopepods)..... id. (,<01>••,d """'00. I(, illa""""'I> ~,7~" of.1I ,he lood ,hey ....R.....I ....01.1<$ ,.....,. ..._ Il.~ mil·1;00 'on, 01 l"ll ... " '.n . nn",lIy b1"'..... <""" ~"ff lly UI ,1Id",,<lIy. In .."''er. ,he b,,'" ."b. , aw".h '0 • d ,., of"'I.id or fi<ll or d.. , n, ig..", notthw.rdMo.' pe ,,~u,o ."..,... foed '"' ~"II . up·pk m<n'ed . in '0"" " ",<0. hy fi,h. Th<I.. n larg . " pengu,n'. Ih. <"'pe'or, . " d,... ~,ng" ,.~c li,o and <qu,d only.

S•••n ' 1" "". ' uf , • • 1, Mr, r",md ;",h . So ulhe,n 0 "• • ,,: (u"' 'ped •• , 'h.e"'''"e... I.ol'''~' Il " '" ,n,1 W.<ldell.'Ie .11110" "' '',lIy cunfin.d ,,, ' b. iceW""" Ww..lell "", I• • '1 ,,,.inly ("h 'lid.qu id , I~~ ","be.'e,·' d ,et " """'" 94~

lIn ll: kr ill ,........ up .11'10 lIt {lie leopard...1', d iet. Th< c·,...,........1. now num_be<illg . boul Ihi" , 01,110"". " ' I>e """".hund..w "",I in the .....Id . The lu, ..,.1,M S"" ih Gwor'" t<etl . In..., e.d..ivd11M' krill. s",,:~, of AntOle"c R. I" , pre·...."OIaI Ahouo n m,IIo"" ...i,h , hioo"""of . _, 7 ," ,lhM I0Il>, ll>e<e ..,I I'Ol"'.Io' ion, annlUllI) <00.11 ''''' m., 1:\0 01,1·

boo ""'" 01 ~" U (Iwo lit Ih"", '''' '''' theeu""n, <"",,, mpllOn ~ y wlL., I«, " . d . t

...." 10 milli"" '''''' of "l" i"Bol«" , or Vl" "~ I"'\ - '«~"5 , w~" I"

of ,he Soo,h. ", Oeea " ;nd UlI< 'h, ~IL""

(in, "'i, minke. hll", pback, " oJ " >1,111",,..,rii lll who,", Belo,. wo,I"" ex plo", .lion. they were prubahl y ."""I (1M" ' Nn..",,,,e "h" ndan' .lId bad • bio'n lI,.Ii""" ~re"!.r Ih.n .. "" "",, The ,....food of Ihe b.l""n wh. le, i. k ll il I"I~. Ih. wb. le b io""" Abou. 45",,1Ii"" ''''''. ond whAk:1 con, ,>«l an eo·'i mAte d 190 mill i"n '00. uf h,1l

Compe"""" to< f'''id ,..ol<.ohl1 Io m".dthei, .iu: "nd thei, nll n""',~ . 8 ) 1971. "'"...h. l.. h.d d..h ll~d '0a biorn, .. 01at>ou , 9 m,lhon 1tM'I>. _ .1>e1.1e . _

43 million """ of k"ll The ,.,.,.,,"" ill,h. .....Ie ........1. .. "" """n, Ih., """ .ISfI,n<lhoo "",, '~ ~nll f" """'11 eo'enb)' ...... Ies ..."" boc"".. . vai l.ble '0 ,he,em.ining wh.I"" ",he , pte<l""I$ , , nd

"''''''''' !>arw:... ,..In ,h. p. 'l 'h i", , . . ... . Ih« . h"

heon ... i""'<He in rt.. pre~"."':y rot. "I'fin ,"d sc , .. ~.I.. " "I ' " 'rroron,. bU L<"" t\lo><!">,"I, de<'. ' se ;n ,h.i, a ~. .. n",L" rlly d ue, p re\l ' I11.bl), ' 0 the ~r..,.,a•• il. bility of food Tttc n"n~e I"'I'" lation m'1 be dou ble whA' i' w"' bef"'.wha l,ng . Th. e","'''" ... . 1""I'<,1."on..Iro ba••••1""""""'" doI;re. ... ,n ,he.~e of .,,".1 m,'"r"y (. y..'" in lhe1950. to 2,5 y.,,~ in .1I< •.,1) 1%0<).. 1.0 'hoo~h1 to he ''''"~_ or.~

lrowlll rAle , based "" food ....._""" .111< 14"-1 'M- joe..... ,••toe P"P"ltl"'"of the Sou,h Groqt .. lur _ I, n ,,,,",u ·. lly hlAih, po"o hably ,.I",ed '" 'be . boll·do""" ofk~ll,

The 'h= ,n...' .OOIld.II ' pen~lIi".,

"'" ohi",,'rnp. Adel",. . ...l,"""~"'"i. ho••shown inc,...... in populo",," Larg' ,n·,-..."', m k'ng penguin< '" thou~h' ", b.<I"" '0 'o<;r f..dill~ 00 " dl... rlnl: ' 4uid,W. dQ not ~now the '"'PU"" 01,1.. r" hand ' 4uid .toc~. 10 t'" ">c" ' '''' ;'' •••iI. bl< h ill, bul ;, i, lik. ly '0 be , ,,n ,I.,Th. ""al qu.n tity o f krill "k.n by .11p'~d",,"" ill ,be SOOlhe," Clroe.n m.y be. bou ' 500 milh"n '0'" pe r y. ..POI",IAI ,oo. and ""tnbe« may <hange .bo' d i,., <' 11 or indi,.c, ly , h~ An'.,c ,Ot:.....ywem depend. on krill. Mu.h ill '.,nokpen<]' on 'he ""mot) prod..., ,,," " I ,heSoul""m 0.• • 0 .._~

.............. ' '''''''' .. 'h~ __ ro ' ''''', L.w~~_04 , -".,.c,.-..:.lI/ S<oo.....O'_.

......"'. 5<......., ''lll~

Th. ~I~,," are ono'her ,m pl"lan' i ro" P of "rg.n;"nsf""lId In ,he pl."~""" R",'lI;ri. a ,. 'h• •mall.OI Ii, l" g ()f

Ilani""" the) .,,, '"""'"""1',", ' ing l. " II, ."d mailY have

fO<ld., Som. I., ll" "'. .. . "'" reIc_ ""'........ "'prod""",,"""II•• thot dli n In the p1""~",,, "nblllley a,. """"" . "'" OJ

"'life "", ", ~ro.....ached 10 ,he..,. ",mom

15.2 Bacteria

,he abiltly ,0 '''I''OO""e by « II di, ;, ;"" "ery few I11"'u l..,.. II<" th.y ",h.bil • r••Il.-.hle . n"ironmen t. AU' Olro ph;cm.ri,,,, ba<i<',i. "", 10 .", tho ph" ,o'1"lh.,izing 9 " 'lUl>.:o,·t••<ill. m",' amonda"l ' I' " " <Tllda l and .,war;"• • ,e a; . nd pro,!oclL'g d. ".. 1>I,, "n. in w. 'm· ....' ., I~ gioo, . " _II .., ill<b""leri••"",i".<1 "';Ih tl", hy<l""henn.1 ¥"','" of lbe dcq>.." fl<,,,, ..h<", 'hoy .,.,he primary prorl"""" (or tt>< ."'....1Oomm ''''' L1~' , ur",,,nd ,ng ,h. vonll (,., f• • 10 ""' pler 17),He,.",tropl"c mar i"" ~••" .... froe h.ing in ",~ ...aI'"

n.e Iypn of plenlu"" "n he <ale~"'" in IIIe k,",donm"phyl•. a"d cI......... ,n It>< fnllow'''1 II,," ,

K"" d."" !II" .." 0 cell, . ,, "' ple .",10"',...- i.i1,.,!;,,1I,le ce ll..... ,.. ,n """"" Of , h" ....A. lIac'", ,,,. •in,1e cell,_ in ch. in. Of gn",p" a"·

IOlropl>,c ","I be...."'0r'"C. ..."ob., . ,,,-, "n.." O·hoc; i"""", . nl " food ""It<e. in ,lero"'f'O!oili"" '

II C,......- blwo.,,,,"" I lg...; ' " '<>1rop1l ie1I"p cell ct..,n, 0 ' I'OII~. I"OO,"", ",meml Ill, '" oea: phylOplriWOl

and ell" on e",,1)' .".. Iable '""""" ,,,,,100 '"1 Ik ocafkx>".duayi.... ",alm,l. Ik lWfoce or 0Ig.n;<m.. a.od p'ece> ofnOll i".~ a"d o,berm.ua,

R 'Cleria pllY . " impOrlan' ' 01 . in ,hi: decay aooh",.~Jow n of or~."ic maUer . rerum;nl il 10 lhe "'••, ba<'"et,.mic.J, . nd O(lInpo und, '" he u=;l ' p 'n lI)' ...w gene,,·I;on' of plants and animal. : ret e, hoct 10"''''pier 14 . Someare al' o able to <kg..d" pullu" ,," "",h", 0<1; "'f'" bad '"eh"pler 12.

They or. , , igllifie 'nL, <IIIli",ortarue maLerial, Ihe 'mall OIL< ot lhe", pll"l",lc•• ,," h ,lie,.an.cl>ed bacle"al ....pul.. ion. mok., lbern ' n Ide. 1food tt><many , m.lI zooplanholl. T1Jey .~ Jmport .nl in ,t>< food

.... " of lbe ne.p ",a , ...he,e 'hey arc con . umed ~~ • 10' B"numoor of 'mall . mmal, .

F'I.lIl un,.i", . illking in'o It>< '\cop .., ha". been a•econl focu. fOl ..mplin••,.;1 ...",e .reb. 113<1., ial -.;n""yon Ihi. m""f1.1 deCle.._ wi'h ,!eplh ..od i. "'''on..1 bek>..2(X(I m (tJ6I.Xl fll. bu' once il ha> .uelled lhe: ... hod 11 ..>Ubjecl'o "i,OIOo. Mc,..ial acti",ly , S.mple, col lec ledfrom 11>0: "'di...... , ..,f""e 4'\1l) m «1 4.llOO f1) deep ,n m,d·<><n<U< ...... of thr """I>o:lo<l"'" "' llanl., """" . fich C""'·",...i'y of .,;1,,,,, """" ",. Il>a, . ....hl< to doc~roJo •..:l l. ....·form .be plan' mOle.i"l f • • le< Ihen had pre " ;".,,I, bc<en_ ....... "'~_" .hmwd .h., ' lor '0""'" ,..... e_,.,. 5 x 10' ""'...;" pe' mdJ.,lde.; n"", u",pl.. ,."".'a,_ 1l-:lO x..... bocl.... PC' m,Il,I, no. dol'''''' "f 11" 0pl.,1 malQ;,,1 slIor1ly "lief Ihr .", ,,,, ph,l<Ipl,"~,on Ill""'"~n.. .... ,mporTanl f""".-rcc for the decp .... ' n,_

",,oj """''''''''''yTho nl",me .n.,,"".......' .... .... deep ... nOOl ",a~

COf"" . ba<.1eri• • ith . bihlla tluoI """Id be "OfJ ...ful lOh"",",,". m 'ara" II pre" '''''I I" ",.re" for ,he.. ,. ~....",... Jal'""'. n",*,", lUOO. 'ho ",,,,Id ·. <lee1'''' ·d,'''''1...- • • l1menfl*. """ be ..... to "" nl Nc~ 1Hl<' ..II,.. ook. pr o ' e . f.o m Ihe bollom of ' he dee p ""eU•.0.,.0' ",i ll hr ' ......frwed In . n "01011'1"..1 I.t>;.....-.y"' he.. lhey u o h< e"llo'ed . tid ,"ol " ed In ,..1<, ond.,

~'" and ""th lloc '""""'" "'mper....... and pH )"'....""",..nI,OI' ""end to ... _ " for ,,"","''" "'''h ,Ila< <. ,be e..M ", J""".'.bfoIc<.·h""l<'f;y ,,,do.I .

15.4

11 Ki""Jo.tI ProI'sJd l"'"'pon' .... """.o«opioc and"'o'lly . int:lc<elkd"'~ aolOlropli> «at,:.. ). 1<1 hctc'Oi,opIu lP'Oi<>b>O)

A , P~"'_ CIuy>t>pJr'f't" ·bf""n al, ..; ,..,1-10" 10 , ,,Iden .....<>Iropbor , ,Ie "dl., in l roopo

01 cIuo.... ; ronuibulc.o deep."'o ""', ....",.; "" ~.

lopI"n l'onI 0"'. IIMtll,moplr.,.:e"" d,.""",.2. Cia ... CIP.,wp/ly£t< rocroI i~> ",Ii·

'''tloo&~II~I''. ....."' n~B<II_,

8 , ,,~W"'" Py" oph",a Ii'" .l~; ..nllk « Iii ...-,thn'1I011o; p""-' ''''" m.... . ed ....... b' ol"",,,,,,,,,",,,,,,con,,,,,,,,. "",.II~ ron.ide,'" phylovlonkl""L C/<u. D",,'pir.W"~ , d"",fl•.t<IIal..,

c , "ItfI- Sarn.b.... ....hol..,..... ("'.....n,k , .n"roopI..kk'"

l> P~"u", Cil ,.".Joor" "'Ii..... roopI. nk'or1II I K'~IJ<,." P/amow pl...... I" ""..,ly """""",Ie. "",I,, ·

,-cllula<. pho!"'yn<br:li< .llol"'fho.... D,..."_P~,,,, "''''''~ aJ. .........,.....

"'0"'1.0"" 0 """",.""", hal><,;" "'" s..rp..o Sea.I\I • ,,,,d<,,,, >4",,,,,,,.. ..., 10; .... llocell"1ar bet·

""'"""," "" Ih 1, «110, " ....... and "'l"".y" ,,,>opt k,.,... For lanFU"'" rwmbc.. of "'"ronpI "'" 101II'IC1'<IpLtnkWOl ), _ .".

~.n~_ I_ '" V,A p~"_ C""'~"", ,,,,, OI C.._" "'LII" ol"'l '

,,,.,, ,,.1 ,,"h,..uc'" and " lIIfIinIcell..I. CIdu H...m..... jeny h... one 1U~ ill Itoc

hre cycle. "",h.d"', IUdI cuIo:>nial f"""" .."""'og""• ......-01•••.

2. Ck".5<JpI><""" jcIlyfi"'.II "~ylu,. C1~"'f'ircJ<~ comb ,.11...; u.fI'lYCmt;

"""'.. .."h CIt..; ....leA biool"",~C P/ryI_ C""~"'~naI"'. anow........ f" >e'

... 'm....n.. u .......,,,,,.. worn..I), "~yi_ 11101/14':'" mollud.l; Il>e ....1·!iU pocro .

pod" plank'MlC,"- Phyi_ ArtI""I>ndiJ ....m.k .."h poIrw1.J""'>cd

'l'f'Cn<Ioge. aIkl hard <KIter ,~cleron.

r, C/<w C,.""""" GOflCpOO' and ...pha.........f , Ph" "", C!tOId",,,, on;", ..I" incl«d,ng "..,.,.

hr.,.. , ...i,1i don;.l'lCf"IIe COfd . tid 1;lIl11ilO ..

""'IC ""110 '"<I<:""lopn....,r, ~o<hJ>IIy/"", U,.,)('''''''''''o: .ad ll<e aodu l" . ,m

....d pule·· Il".ac. ... Ip'V, lII~fol'/,,~kl(>'" 1.", ,,1f""", f."", "'" pIIyl. Anne l'da

(.., gmrn Lcd "''''''', ). M" llu""a ("'<:Ilfi'" "tid . na,l.),1\, ' 1""1>0<1' (et .1" . "d b.llac le» . ('.ch;"odcm...,.(,"''''''' and sea urdl,,"l. ' "d O1,.-d. .. (fi"'). Sec, 1"" tn. cb,,, fl, auulJ, u",mOl 'co of lhe: ","~IOO ("b.r~

Ie. 11» a"d I h~ benlhos (In ' hapler (7).

Tb< biolog;".l ",~.nogr'phe:, ne"d' '0 I<J)"'w wh.' "I""', e.of plan" and olllm"l, m.lk" up Lh~ pl' " kL"" '" a gi""" ge<>g'.ph;" art o. Ihr 000""",,,," of tn."" o,gam""". and ,.1,...

J~1: Sampling the Plankton

Cla"sil'ication Summarynfille Plankton

15.3

Viruses in the Quam

A vi,,,. i• • ",,,,,,,,110101 I'"I100Ie lJIOd< "I'of ~....,'" "'"' 1 """co:Ied by. pro."',n co.l . V"" ~ig~ly ,,,,,,,,,,,ful1""" ';" ' : ,I><, ,nf"'1 pion'. '''i'''''l••DdN.:l<r,.1 cell•. V,ru .........._lOe,,,I)'on ....... bol,. OI:1"" In by 'I><m>cl.-e<_a n ",pl"." ,1><",,,,,1. <-> onl, ,,,,,d••ho" • • ,.. l)ul(oll •• pli""'",o ,h••itu'

,,'", ""'l~ of ,II< ~"",'. ~"""': i'"'"' ,hen lC P"" Iht;e groe. 10 • n<wu ll. _,., '" .""hans< uf ton<be ""'.,i.1 bt,""n OI!lOI'''....

R...."'ly. microbiolo~im ~.".. disco.ered . " "nup««d "o ,,"ol.lKc of'i,,,,O$ ,n m",", w. ,,,. ( 10'- 10',i,.. 'n/ml). 0"• •• " " o f .. mpl••, "" <1 e""". nrr,,,oll,"1 , in" .. be·' ~ " i ~ o"d IS ~ 10' p.' mll h l,terd"f1 "~ (II< , "" i"~.0<1 ' "'nme' b'" d"'l'l'ilLK ,,, Ie" ,~ , n W· """,c' po< ,,,ilillil"in The win '" M,, " of ' h, •• 'u ••, 01"p<O,o; l 1<, I>< t,~ ,. 'Oe wate , . bu' ''''''.........._ ....ed i,h Nclel". I' h.. been

."u........,. 'hOi hn<te"" fl'''''''''''''''

is main,.in<d '" balance by the II""'" .,.-"n,le<clfed _,,,"<>pi "", the"" h"hcOfI<.e.nra,iOfl' 0( ..." ,ndi..oo ,.... ,....I."r........ ,,'",be.o i, ,"..' f",lOr

•• tI.. ""oI",i,,1 COIl' t'<Il "I plont10''''b.M"le," ond Ih.1 . " "i8h' O<1i""l,. "·b.1<ll!c g<toe, bt, """ """"Ioto""siofc<1ed b, the " 'n<>".1 ....."

OIlier " ...."'he.. "'1"'" l"lh ....Iobundantt io ocean ....... >rod . 1", 1, '1.n"",bo" of b...·...i. ",10<'«1 w,'h ,,,""",Up '0 1.. of 'hc M lorut"'p hic hoc'c"' .. tId 5%of It>c cy."oNc'le' i. /""1' ,I,v. =m. , in. I~ O ut i "" . <u,,,.,,••ol "' . ,,".' im"" Estll". t•• ,~. ' n,.1 .." I i ,,~ tloo

f ro m 1111. da" cond."" !hat . br.., 32% 01' he hotcrolruphic b••:lo"'a . nd .1""" 15'1>of lhe cy.nobact.". <ool.in ,,,,,"," """P""rle<'f an y It""" """' . inrl,c "iR~og.,n tl1IO' 'i"l ,,,r...,;, ... ""'y Ile a "8n,t;<001 ......,h.n"'" of ",o...h" in marllleback"O

In l.b",ot<>JY «PC"""''''' Ille add ,·tlnR Ot ..".1 1""';01•• l'edoced pn"",',

pr<><b;'i.i', b, It> ....o:h '" 1M iod",.. ·iOIl ,"' , i"l..bon by v.",,,,. < kt he afoe'o, ,,~u1,"ing p1tyt<>pl.."~I,", ..""mo·

nIt~ "l\lCturo and 1''''''"'' prod,.. 'ivi'y ,I'

,he oce.n,. Ileca""" .i",... <an ,"foct a,",'."y 01 mar",e pl>ytopl""oo, inc lo>ol·'"8 'h ""'"" tOeY m.a~ . If"" oc.... lood"'.'" by ",,,,l<h 08 0' l. rmm, ""~ I'lty'o,pl, " klOn bloom,.

T he . bLl lld. "co of , i.." o, ,n "C<an.... len; ind i"",es ro"',,,e .. r.i JIIfO<' l i (~ , of' qu, tic oocteri', and Ihi, i, Iikoly ttl mo' "If"t nOIO..1go"elt< on~",«", ni o'pc'i ·tn.",,, in I=<o,io i p<'pul.. i<In' hOY.~"

OC'<U'f1ng for 0 ".!)I lo"g "0'• . Ne w teo

«~ITh ,-m., the I"",iblh'y 'hal ~,a,i ...b.cte",. ,i••" . 1 o,ehanco o r ,,"" ..m......1. mig hl bo . 1>10 '" de.eloIJ mil"

,••H'. 'n .,,"h;otic5 ""'" ,. ;oq_u1,...,.

o.-.~ llC<l"'''''~'' ''''''' ....r"".lly O'~i_ml bac,.,io ,.roo..-.l ,nl<' ..,.,,,.1w.....

,n ,he w.la col"""" ,hey .... nal"'. T'adthOO.f1y, pl.nkton'n: ...mplod by ...... ,n~ rone·,ne<f1. con.·""'-ped ,teU (f'l

I~. f7) 'h""'4:h ,he tea """ond .....d Of by droppin8 ' "'"",,,_III down .,...' 'he ,i,l. or • n""mo~,ng •• , ..1 , ndpurl,ng ,I ba<k up h10 • bud",. AI", .. ,he "'" is "" wn.:d '0lhe d.d il is ,"""'" core/oUy. lind II>< "ca",h" ;, collected ina 1.1",led J"'. If 1M 11<1 h"" !>cen h."I'" vert ically th roughlbo ...... . in""" placc, It>< "o lum. of wale' that h., f'l<''''~

' 1',"1I~h 11M: ,..I can be ""kul..ro frum tloc aren of Lho "", ',mouth . nd the d"t,"",. t~""'fh w~ lC h the ".t w"'pulled. II',"'= no' "' .. 'o",od beh i,~ 1 lh< '","i, the ,'oi" ",< e.n I>< c.l ·cul".d from II.. !,me of Ih. tOW .".1 t~e <peed of th. (Ow,",h i"h "'il l I.U Ill• .1"''' '1<,. (o w.ol. Thi' inform.tion c.n' hen t>< u'"<l w'th 100 .re. "f the ne1 open'ng to f.nd til<: >01·"me of W"cr thol~ , t,,-r... gh the ne1. HO....,>ef.1O me.

,u,. Lh" w• •" ""I",..,h""I,.ocI ...".-I~,. fl"w ,nr't<.i . plac'" in Ihe " .....'h of "'" ne~ The eatth is rllen known"" 'he " • •1~,~""'" of ....... """'pl•. oot ,here i, no ind"", ·' ion of huw lhe . pecic_ a,o <li"'-;f><ote<l within ' IIaI . 04urn<

11 " .1.., po•••bl< '0 ~i"" .nd 10.. .,.- ,he nt:1 "" • i.re.", 10...-..1 ho"torurly. nu. """""""" ..1.......ampl,,,,10 be av....,... OW( botb d ond defl'h . II Ihe ...mpk;is I<> be I. k opec;f.. dol ,he .... 0'0' re low.r«!olosed and " op<lltd w",,", lbe de;ired okplb i, ",ached

Af'er lbe IOw,n~ ope'''K>II. the ""' " "il"'. d",,"d belote itis b",..,h' "" Ib'<K11f' lhe >hoUow•• "'01..

Tod.y. <>eo.nog' .pil." .,"'n pling lO0 l' l. llkto n a'.m..luple · ...l '1"<m, n"","led on .. ", . g l. fMrlO' Ihe "<t,••• opcoorl .nd eh,..d o n com m. nol fTOm 'he ship. The

Figu,", 15.11~ "..,,,,,,, "" ,,_.... ".' • , ,,, ,,~..-f,,,,,, "'~'.

T", ic bloom. of so",Ie-<:<llod OIgon" ,n, Ih. 1 discolor ... ...."" 01' oft," loo..n <IS red lid,.. : ""'Sl " .., produced by",rto,,, 'f"'d coof dino n ' l"'I1.,"" n.... h'...-n' m. y." m.," ,, ' b< !XI''''"''''' '0 fl<h . nO "'her or~,n"m.< a,," m.~ 0 'moY !lO1 p'od uce ' Y" LI"" " " ul l",,.ly'i, "'d lflSll l",i,,,,, iHJ(!>~Pl. 1"' '' ' 0<0"' , ' bolli'''' 1""""'1li (NSP). <>< d,.' nh",i<

f,.me .!so c.",e, .kct"''';o ,en..,.-, ' llal ,e l. y do'. ""..~"il )'. ,",,,p,,,",u,~. ""te, n",,·. l'gto 1c:~e1. ''''' def"h••nd" III< angle 10 ,I>.; .lltp" comruler,

Pla"b:m low, 1'1".1 be l'lIpod ......~. ' 0 ca,rnl"" ", .ll''''''m. buo .low mooll~ on It, ,..., "'."', I"'" lhrouaJt ,I><...,. If 'ho ",.. " '00 I.." ,"" ....... "'ill be f""W .... . ~

f''''" ,"" """,th of .... " . ' . " HI I• •• "' ..'" ,h. " " 'p<c 'ed "" II"" ftll.rOO. SIZ< of ' '''' ",,,h It'OftI "'hlCh 'he ocL j . ", ..",also ,nil"",,". III< c.leh; a Itne·m"h nol clog, rapl~ly ""d• lo,ge-mesh '>el l o~. " ' I,,,,i,m . ,

PI ." ~ , o,, n>Oj ' al,o .... f,I.. ,,,,o fm m , . mple. "k"" hy"'at e< bonlc "' .. , ub"",,,,~I. PO''''P, W""therlh...mple i'",k... ~~ .... . ho'"e.... pU"'P. 'he ,"'" "CJ> i> to dote ...,i..,be ""mher ..HI kind. or pl""k1on ,n , he <ample 8<:<.•....,II>< ,.."ntoc" of ............ . re ... 13lF. in m"" <.... i' ..... "",..hIe 'n d;,«U~ •• """,1 '" «III,,' ''''' '''' 01 ca,<h,I..,... "'" CIO"~ "'''y bt ",...""Ir ....bdi.Od«i . and a ",I>..mple m. y be c h«~"d undu 'h" micr o, <op e. If ,h.am"u,,' 01 p1,nktoo i . of ,rut.. """.a t tha.. lilt ",,,,j, ~f

...-g. "i,m" , n do"""i"" ll>e q.aOl" y of >uU.

I~." k '"" dire...ly, the e•.-I", ,e,",ned <0 ' l>e sJ,ip i, ", Io'ed 10,he <len"'y of tbe 'OOJ.Il,nklon b'" docs fHl t detenn".. ,0<'f'O<' '''' pre<.enl. Tbe . bund..><oc of ph)"oplank' oo COl' be"",,,,"',,lOd br di ,,",., lhe chImOl"') 1I pi~m 0111 oftf>oo ...mple.OO ""' 11$1.... ,"P""'" c""""'""'.~,ooo.. rio","1""'01 i"'trum"",.. i<H1 hO' l>c<n don,~ned lO "",a"" ,. 11>0""araj lluor="a1l ,if"al , n,,'ll from <hlorop/>yll I"~n..,,',n f'lIW>pl on k'"" ,dl •. T ",.." ".me n" 're mad• • ,soa, duecl ly artd ,"".n""e"",ly " deplh, and ' he" rel:",,1", f'lIyLopl," ,'o n " ""ndo,,,,.,

!I II ,~C'" me' hod, ,'11" .. ' 0;' ''' ' ''' '" re<or<l popul• .",," ,lol,. iLi" , '" 10 in'l""" i" d " ,d" . 1 o r~a"i ,m '. bul lh<y00 nol lell u, 1>".... [he" OI~;""""$ hc~.ve i" thei, awin..•01<0', A no " .idw pl." k'on fe~Orde' d"" Sned . ' tbeWoods Hole O<.£aoov..p.ic I""i""."" """•• strok li. hl...i,h rOll' .;.tro "*'""..,,"' I"''' d ,m-rml m>t~.r",",inru tnI"""o~aph .... pl.n~""'. "Tho .. rob< l1....... "x'y '~"'" 0_<H><I. <aplun ng in,,~e. of ,I.. ,.-~...i"'L> "'" ",.,u,<hcuhnpe ""II .11""," ,be,n I~ I...m abOll, 'be ,..- i",mi nt , leed", •.arHI rep}(,d""liOll nf ,he pla"" o" . Evem", l)y ,t><y h<1pe 10prngrml l!>o .y'''"' 10 ,«"~,, ilO ,lo ff",enl linds of pla"~ '

10" , . n, blillg ' hem '0 "" I a,,~ ' I'<'<i"• •",1pop" I.' i,, " In f.,, ·Tn ;"'"" w hile Lh. ' y"om " f" "",n~ al "". ,

15.5 Practical Considerations:Marine Toxins

&h<llfi'Q PO''''''' ''I: IOSl') ,~ l>um.... """J er..".. mo ......,.rodoy".'" 'b.....~ .'g<>led ,100 d,~nall"I" .... )n Non~

..."""",on '". "'" S<'o"<'<lII d,lle ....." "i"ofl~,,<lI"'" prod....,

..... hd e.: Gow!""",, . M.."n d" ..... •nd Gy~",odilfi"M

(pn:.ion>ly P,.,..lttuJiJ,·.,J) . ,. 'o",c , bul II''''/I!~cu" on'S"" r, ~"re 15.6. Sl"'ci.. " f """ym,l". ' nd >lI~ ' Qn<It , ,, ,,,

loon,l in Lempe,"'e l.h'U"'" ..~ l!"n.'.I1 ~ nonlOX'C ,,, ,r."<!>o lin", rh<m, d ' e" I"" Ihe ,,,, ill' I'"xlll<-od by Irw. . " ,.non.gell.le, .1. C(\Il'''O,,....~ '" Ihe 11""0' of Iho ,bo llfi. h• • lhey roed on the bloom, 11 " the ,,,,;n, ' h' l prt:H.1= PSI',!'I'P. ,.- D.'iP, in It>< h"",.n. ,.., n~ ,I>< ,fleeted ""'rtf,,",,

G,,,,,,,,odini.,,, >od • ' I.... i.. of """....,,',,-' fou nd in '''''",,,,,,,,, ~a'o" of ,"" Gulf of ~,,<:<> k,11 fim. G<m",oJ",'.100 blI••hn",1' . <HI (;nOb; G_ ..bm"110 oJoe" " ,,",

It"l'f<"''' '0 _ -=......... til.. , lie "'" lid< problem.. ~""'''' i .rwnd 1100 "",,\I, In 19~. majOr 0",_ <tl<ed 'idoe <ou,....{ hy Ihe d,,~'floi<'II. 'e Gy",n",};ni~m f>,.ty,

.",,<k ,I>< n ,., Gu'l Cu..' boo",.-ho. , I' ' preao ';OO ~m O(Xl",,1<., ) . lung ,1>0 co" L... ! ~i ll<d mOle ' h"" 22 mjl ~on f i" ,o' er more than ' '" 0 n" " " h" tt. ....."og 01 -,hellf"h w"'h. "" ed along 'h r.~ ·4tia r l." ut" 1>< 'k<a' coasl so" , h ofa.h,.""" 111< <l.le of Tc... lu,' 011. " 1 $I.( m;lIi,., 0'O},,'" r>rodue',on, ...it!> • "~,,I '"","'.... ie 1M. nf $' 7 mil·lion, bu.""",,,,, based . w< ' .... ,i.'" ..HI KCrea'ion .01(..-.4..."",Iy. 'Thot """II: ye,,, h","',e,h'" bol,le·~ool l-tlin."oed . 1""& II>< ~oas" of " '''' . ....y , .... M" yIOnd . Ab loom of G. "ro.; <>f( u" . d ,n ,h . "'..m "" Ie " ofrFlonda ', we" "",-". "The "in',,' ...., ""I": ' h< red lode ,,,,.vi.cd ,r.: "in"", . .... "'e """...nr <",OcoJ II .""a><l FI,~ "I "

a,HI up ,!>o E"" ("<)0.'1 of ,r.: lln ,tod S '" t~. The red ',do ...·,i • •d off 'he Cm, li".. "' . h" u' ,11, ,.n", ' ime ' he '"< Ll 'h" I, ,, f" h bega" Ihe" . ""u,1 , ,,mOle, O1 igmlio" up IheE,<t C""". T!>o O1e"h",l<" fed ,",' the (; h,.,v; , ,,d ",cum. ·I "o~ lhe ' 0<1" ill 'h<i, li••,~ An", ' II<' dolr llln' r.... on 'h,rncnh,ik" , 'h< ' 0' " ..,""~... Ihe 10<"'1 01"",1", of the dol·ph"" 0' c , "1 ,he ''' '0>01> ' 0 lox !>o., """on'roIl"ly,loll,n~ ,"" In 1\ll!7 ond I ~~~ . red 'ide >""'ll .to: o"Ire-. of Florida "P'" "",-.ad """h",.td. thi' time ' 0 """hCa,oli... and api" d:lfpll".. d'ed 1hc ,h< Hfi~ ...ellloIo..d >nd <C<J<lOtrli< leo"." ..,ach«l $2~ m,llion In 1990 the

f"" wof"n'l«:! ou'I'n,.~ of 0$1' 'I'",,,,,,,d '" N""h An..,"'"and w"" " a<e<I '~ 0 d" • .na~ell.l< bloom;" C." ",lj. n "" 'e<.In W9l PSI' 10Xln "'" ,10 1<e'eJ f,~ the rh'l lim, in Il,e It""or Dt'tllle""" cmb, (' <N n !II... , " ," 'l<'~,

N" o ne kI,o"'. pl <o " d y ...h., "i ~g<r ' Ihe ' ''O"enbloom of 'h<oo Of~""""'. ho,' ,<d 'idr"' "lten h' l'p"n in>fl'Ii_1t and IOn''''''' .ft", I..,.. y <~... h.ve prod,occd a land,.""rr of nu'rio"'ncb w.«,. o.""na"e W."" "",,,lIu •cy><<If """n~ cdl ' bol 1... 1<> ""uom and ,n" ,,~ "'"oed""""",. In the !low 01 Ixoy<.nd ~..to ,,,.. '4<" F...glaod en'''' . " ,nl" " O"'''' ,r>d ligh' ' Ppe>' 10pi..) ",ajOr ,01.. ;n ""t i• • "n~ 'he " " " . In ""'p'" c" " ,. ,.... ,. ,~ 'Ile ey'" ",em ,,, h.",. """,al bolog;c. l clod Of

",,",.Jcy<1e of f" C" ' ''' ;C41. I' " U'ool ll' U"" ,,,,k,, n d,,·lurb,,",'''' 01' ' he bo"<:rn by <llbol n"' ",,1 or . nilic",1 ,,>eM.(, ,,eh "' . lu",pon g of , edin.." ", 0 ' d..<d~ ,nil " imul" e ,he<y, ts i",o "" ,i. t'y ...,d '''I'"t f1" r"~" ",, ,,'t

Some scientists think that the continuous addition ofnitrates and phosphates to coastal waters from bothsewage and agricultural runoff may be partly to blame forthe apparent increase in the number and severity of redtide episodes. Extra nutrients mean extra growth and mayexplain why instead of a series of small blooms, theworld's coastal areas are experiencing such massive outbreaks. An additional factor may be the increased commercial ship traffic that brings hitchhiking cells to newenvironments. In a survey of cargo vessels enteringAustralian ports, 40% were found to have viable dinoflagellate cysts in their ballast tanks. Other scientists do notbelieve that there is a rise in toxic blooms; they point tobetter statistics, communication, and reporting as well asincreased eating of shellfish and fish. However, the experience of the Japanese in the inland sea of Japan suggestsotherwise: Red tides increased from forty a year in 1965to more fhan three hundred a year in 1973. In 1972 authoritics introduced controls to cut the nutrients enteringthe sea by half; the frequency of red tides peaked in ] 975and has been decl ining since.

The toxins that produce PSP are powerful nerve poisons that can cause paralysis and death if the breathing centers are affected. Some species of red tide dinoflagellatesproduce several toxins; one species of Gymnodinium canproduce at least five different toxins. The saxitoxins thatare found in butter clams but arc produced by Alexandriumand Gonyaulax are fifty times more lethal than strychnine.The toxins are not affected by heat, so cooking the shellfishdoes not neutralize the poison. Even after the visible signsof red water due to a dinot1agellate bloom have disappeared, the shellfish can retain the toxin in their tissues forlong periods, and so the beaches are kept closed to shellfishharvesting. NSP symptoms are similar to those of basicfood poisoning; airborne. wind-spread cells may cause irritated eyes, runny noses, and persistent cough. DSP outbreaks may have been occurring throughout history andbeen attributed to bacterial contamination; only recentlyhave researchers linked DSP outbreaks to the presence of ac1inotlagellate. Between 1976 and 19R2 more than thirteenhundred documented cases of DSP occurred in Japan; in1991 over five thousand cases were reported from Spain.Other outbreaks have occurred in Norway and Sweden andalong the French coast. Scientists speculating why the toxinis produced believe it is a defense against predators.

Just as not all red tides are toxic, not all red water iscaused by dinoflagellates. The Red Sea received its namebecause of dense blooms of nontoxic cyanobacteria withlarge amounts of red pigment. The Gulf of California hasbeen called the Vermilion Sea for the same reason. TheIndian Ocean has red tides due to the presence of a tox iccyanobacterium, not a dinoflagellate.

Other Toxic BloomsIn 1987, at Prince Edward Island, Canada, three people diedand more than one hundred others became sick from eatingshellfish contaminated with domoic acid. The domoic acid

386 The Plankton: Drifters ofthe Ope1J Ocean

(a)

(b)

Figure 15.18Electron photomicrographs of two xpecies of the domoic acid producingdiatom Pseudonitzschia. (a) P./ulJlg('l/!; (lOOOX) responsible for amnesicshellfish poisoning in PIince Edward Island, Canada. (b) P. australis(l400X) closed U.S. West Coast shellfish and crab fisheries.

was traced to a bloom of Psc udonitzschia pung ens ,(fig. 15.18a)-a diatom, not a dinoflagellate; diatoms hadnot been known to produce toxins until this outbreakoccurred. In 1991 pelicans eating anchovies off theCalifornia coast were dying from domoic acid produced byanother species, P. australis (fig. 15.1 Sb), Shellfish andcrab fisheries along the California, Oregon, andWashington coasts were closed.

Domoic acid poisoning in humans may cause shortterm memory loss and is called amnesic shellfish poisoning(ASP). Other symptoms include nausea, muscle weakness,disorientation, and organ failure.

The reasons for the sudden appearance of toxic levelsof domoic acid are unknown. although speculations aresimilar to those proposed for the dinoflagellate outbreaks.One researcher has suggested that when excess phosphorusand nitrogen are present the diatom populations grow explosively, and when the phosphorus is depleted some

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14 Ef>tSO<ie> oll<>.<lC pbysoplaolklOn l>kxlrnr. _ 10be

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'ktfyr,,,,, _"'""""', .

the plankton: drifters of the open ocean · figure 15.7 zooplankton sample (enlarged)....

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