HomeMy WebLinkAboutAPA1962Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
EFFECTS OF TEMPERATURE UPON YOUNG CHINOOK SALMONSeymour, Allyn HenryProQuest Dissertations and Theses; 1956; ProQuest Dissertations & Theses (PQDT)pg. n/a
EFfECTS OF TF~SRATURE UPON YOUNG
CHINOOK .3A.LiiAON
by
~LLYN HE~~y SEYMOUR
.a. thesis submJ.ttod in partial fulfUlmei.t uf tne
requirem4nte for toe de~1"'f.l.e c f
DUCTl. rt CF :-:-tlWSCt HY
U11IVE.1..SITY OF ~'/ASHlNGTON
1956
Ap)rovecl by~~~
Department _c2"""-~~~.:.:;;;..;;_._' ~---
~,2> 05-~
l.'!':I\TRSITY 01' \'i'ASHJNGTON
f);otc: Jcly 19. 1956
We h .. , .. •:.an·tull~ tt·;od the tlu._;, cntitkd E!'fee"\.s of Te:::pe:-3.._u:::-e
,. -· -· . . s·-·. ~·?-:):'1 .:.:r-1:.: _ vf'U.nOOt: .;.u.."'!\\n 'Uinuiucd 1.,·
;.llyn :ieo:l!';.• Seyno~ in p;artial fulfilln~t ul
De·~C o~ Doc ... or of Phil~sophy dlf' rt"t)UiJ~u~nb ul the
and rt"Cun1nu·ntJ ih acH'f"l.otnct". In ~uppou ul thi"'" lt"1.urnntt·t~tlation 1\C l"C"!tCttl the
fttil<J\'l ing joint ~tittc:uK""nt oi (''\aluoulon to he flln.l "·ith the thn.Ut.
!·::. s~~'.lr ~.as studied ti~e ef!"ect.s ~r .... 3ter t.e.::;>erat.u:::-e
~?O:t t::c e"L-1:~"' cc·:~lop~:r:. :.·~ t.!:c ch:.!1o':"k s::U..non. t:e r.as
a::r::~o:-t:~cd t:1e ~!"'oble:n z·:.:-st. 0:1 <:vti:.:3.t.!r.-: t.Ge ei!'"e:ts o£
tii:"':""e:-c!'lt t.e:-:nc:-at'::es ~el~ c;:,n~:.:l!lt L'-:o·.:~ho:Jt ti:e -;"a:riod
of :..::·."'C~~i--a"!.:..on. A se::o~d scr~~~ o~ eTa?~!'"i: -~!lts ve:-e nm
o~-~~r:r.--:..·.e e:--s !or t-a....,·::.n~ !e:1-:.Z"".s o:'"' "! . .:_~at e:.ty
•:t..._e~ t.e=:;:>e:-a"t.::rcS ~!: :i ·.~.en ci::'\r.~i.!:-t.:-.e;, :o ~~=-= c....:s constant
te.~:-:-::~:::"'cs f'o:-t.:-,e rc:-.~:~::c~ =-~ ~ _,,.. tine. ~~ ~-?rird ~~~ies
«3~ :::-c-.!"e:i <-t. "-a:-:.::f~lc t.e::~.-:-:;-.."':-es. In this l:1s't se:::-ies~
e--s a'-":"e obt.<l'i.!:C~ ;:'ro:-: t.nc S~:a-:i.t. '1:::... ~e:1 !-•. cvers in
~:est.e:n "t~":is:~::: -:.:ln, t.he i:.n'":.i:J..t ;::i ·.;c:-in L\lst..ern ~:as!"'.i.~~
a:1:i !-:-:::.:-:. -:.~e S:iera.-.ento ?..!. vc·r ir. :.:U.; .:'"'o:-nia in o:-cier to
e~...l!.:~:.e :-:'l~ir:..l :ii.!. • .!"ere:~ces in te:~::,c:-:l!.t:re ef~ect.s. Control
lots ~e:-e r~ in all cxpc~-=cnt.s. ·
'"::u-o-.;..-h t.ilf=!Se eX'!'!':--1.:--:c~ts !::. Se:.~cr..:r !::1:: C-O-r!!'~-ed the
di!":'e:""C-nt :-:Pt:-~:>ds of" mea,:: .. _::-:: nt; te::;~crat.~e e!fec:t.s ~--:d has
ce~.,nstrat.cd a ci:Te::-enee :Jct.u~e:l r.ces :.n the e!'.feet. o£
t.e=-.?:-a~::re on rr,-:,e or develop:"lc:-.-:. ::.: -...ell as :nE c!'!'ect.s or
t,e:-;:,n-~·-~ 0:1 d .. -at.i,n 0;-. ~t.---:n,--c-.;od ,.,~-·-'.;t-· ra•-• •• ---~ ..... .. ·--·· . -.__ ,. .. .., ... \ooc.L,L-•"' ~.
abr.~~~li~ies, ,-ro·Jtt ~d nc~is~ic c~~:"'a=~~s.
:·:r • Sc:::-:to::r :-:as de::tOn~t.r:lt.cd t.is nbili t.7 T.O C:r.lcbet
oriri:!::U expe:-i::cnt.al. vo:-k ~'-'"!~ :.o ml·'ll:-;:e a:;,j ev:Uuau the
:-es·1l :.c. :::oth !lis exoerine:;t-:ll Oes"! ~ ::...~d execution are
ext-elie~:t ~:;ci his theSis is worti:~ .. -:Jt accept.a.~ce as a
Do:to~al cii~=e~tation.
J
TJ IJ::SIS R£.\UIX(~ C0.\1.\1 nTll: ! -. ...... ··········----.. ·:-t" .... ~.;..L.t ___ •...•••
,cd~ c. &...t: 7
. -""":'_.., . . .. ···c?:.·· . .. . .. . ............. ..
-'/Ct. --.~.-·~ . 'A..-1'
··•••• " · • ··-·· " · ••·•· •'-•······•·••··•• •·••-•••••u·-••••u•
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
In presenting this thesis in partial fulfillment of the require-
ments for an advanced degree at the University of Washington I agree
tnet the Library shall make it freely availab]_e for inspection. I
fT!:·ther agree that permission for extensive copying cf +~his thesiz fer
schclarly _purposes may be granted by my w1.jor professo~: or, in his
absence, by the Director of Libraries. It is understood that any
copyinc; or :f=Ublic::;.tion -:JJ.~ this thesis for fi:1ancial gain shall net be
o.lloHed Hit~1out rr.y i-l.Citten permission.
/c~/.. .. A._ ;4/ ~. ~ Si[nature ~ ~
Date ________ ~_-~~· ~~--/_9~~~£=-----
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
ACKNCWIEIXlMENTS
The counsel Lnd help given during the course of these experiments are
grate~ ackn011ledged.
The experiments were conducted under the guidance of Dr. L. R. Donaldson,
chairman of the Supervisory and Readin.:~ Comad ttees . Errors and caiseions in
the manuscript have been correc~~ from suggestions made by Dr. R. Van Cleve
and Dr. A. C. DeLacy, members of the Reading Com:rni t tee. Valuable aid as to
techniques and :methods wu received from Dr. A. D. Welander. Calculation of
the t.hreahol.d temperature and 1 ts confidence interval was suggested by
Dr. D. G. Chapman and r<r. c. o. Junge.
Ch:1.nook salmon eggs were obtained .fran the Washington State Department
of Fisheries through the cooperation of ~fr. C. H. Ellis, Mr. S. Fallert and
Mr. E. Fisher --and fran the U . S. Fish and Wildllfe Serv.l.oe with assistance
from Mr. c. Atkinson, Mr. J. Pelnar and Hr. R. Burrowa. Mr. BurrOW8 ale~o
made available same of his unpublished data. Mr. r. R. Olson and !1r.
J. R. Donaldson occuianally assisted in caring for the lots. Dr. R. Rucker
and Dr. E. ·Ordal made pathological exam1natione of the fry of Experiment III.
The technique o.f radiographing small fish was lea.:.--ned fr001 Dr. K. Bonham.
This research hs.s ~een supported by the Applied Fisheries Laboratory,
which is under contract to the u.s. Atomic Energy Commission.
The wOl"k of cheeld.ng and typing the manuscript and bibliograp:t;:· was
shared by Hiss H. Chase, Miss G. Brewer, Mrs. M. Pinyan and Miss M. Johnson.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
TABlE OF CONTENTS
Pa8e
I. INTRODUCTION 1
II. PIAN OF EXPERJ:MEN'l'S 7
Exper.!Jient I 9
E:J;pel'imant II 13
bperimel'lt III 14
nr. MATERIALS 21
IV. METHODS 25
v. ENVDWNMENTAL CONDITIONS 34
VI. DISCUSSION AND RESULTS 38
Rate o£ Embryological DevelopDBnt 38
General Expreaeicm.s 38
CO!Uit&nt Temperature Experilnents 47
Altered Temperature Experiments 65
All Experiments C<Dhined 76
Mortal.i ties 84
.Abnormalities 89
Growth 90
Meristic Characten 96
vn. SUMMARY 114
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
F~:ure
L
4.
7.
8,
9.
10.
12.
13.
LIS!' Ol FIGURES
Average Water Temperaturee by \'lieeks at the
University o! Waahington Hatchery,
Nonni>er 1950 to July 1953
Chinook Salmon, Pa.renta o! the Experiment I
Egg l.Qts
Lot Temperatures for E.xperimeot I
Obeerved Temperature• by Day-a for Lot• of
Experiment II
Predicted Tem~~ratures from the Beginni.ng o!
the Experiment to the Fingerling ~~a6• for
the Lots of Kxper~nt-III
The Controlled-tem~erature Hatchery Room,
Fisheriee Center, University of Washington
The Average Temperature and Number of Day-• to
50 Per Cent &tch for All Lots Reared at
Constant Tem~~r&turee
T.he Average Tem_t;erature and Rate of Deve.lotment
for Egga of "t.he Chinook Salmon from. Green
River for E~riment e I, Il and III
The Relationship of Temperature to the Logarithm
of 1000 / Nwut>er of Da,y& to Hatching
Three li:atimate.s of the Threehold. Temt=erature,
a, for Eight Lota of EJq..eriments I and II
The Helationahi,~--of the I:.lelehr~k and Loginic
Equa tiona to t ha Rate o! Developuent Data !or
Eight Lota o! E.xperimenta I and ll
The Rel.&tionahip o! Tem}.·erature and the Number
o! Daya to Hatchin;), to the Logi.tic Curve and
Ita Reciprocal for E~.ht Lots of .&xperiment•
I and II
The Average Temperature and Rate of DevelopJlClt
for the Egge from. Four 'tacea of Chinook Salmon
in Exferilllent Ill
Page
6
10
12
15
19
22
50
51
56
59
62
64
69
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Li.t of Figures
(continued)
18.
l:t.
20.
21.
2.2.
2.3.
25.
26.
Z7.
Cumulative Percentage of the Number of Egg1
Hatched by the Mwnber of Daya Arter the
Start of Exper~nt III for Fcur Races c.!
Chinook Sal.,a)n
Rate of Developant anQ Temperature for All
Lots, All Exferi.Mnta
Relationsbi_ti of the Belehrade.k: Equation and
the Log iatic Equation to the Rate of
Development Data from~lll Lot•
The Relationship of T~erature and the ~umb~r
of Daye to Hat.chl.ng to the Logi.et i.e Curve
and Its Reciprocal, for All Lota
The 5 to 95 Percentile ~e !or the Hatching
Period of Chinook Salmon
Cumulative ~rt.alit.y in Per Cent by 'Neeka
for lots Reared J.t Conatant Temper&tures
in EJqJe r i1Dl'J nt a I and ll
Temperature and Per Cent Mortality for Lot•
Reared at C onatan t Te~~perat ure e
\ieight C~ves for Ex}:;eri..ment I
Radiograph of a Chinook Salmon Fingerling
.kveraga Number of Vertebrae and Temperature•
for Lots o.f Experiment a I and ll
Average Nl.IJll)er of Vertebrae and Te.mparature1
for Lota of Exper.iment III
Temperature and Per Cent of Chinook S&lmn with
Abnormal Vert.brae for Lota of ExperU.nta I,
ll and III
ATerage Number o! Dorsal Rays ani Tempu-a.ture
for l.Dts of E.xperi.mente I, II and Ill
Average Number of Anal Ray• and Temperatures
for Lots of EJqA rimante I, II and III
Page
70
78
80
Sl
83
85
87
95
99
101
103
107
llO
ll.3
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Table
l.
2.
3.
5.
6.
7.
8.
9.
10.
ll.
12.
13.
14.
15.
LIST OF TABlES
Tempi'raturea of Some Pacific Coast Rivers at the
Time of .:) pawning of til e Chinook Sa.llwn
Sunmary of Factors Related to Ex.t-Griments I, ll
and III
Measurement Data of the P&rente of the Lote for
Ex}er iJnent s I and II
Mineral Analysis of City and Well Water
Compari5oo of the Daily T~1J:i.lera ture Range for the
Green and :3ac.romento hivers and for Exper i.mGnta
II and III
C:xye:;en Content of Water in Tanks and Troughs
A Comparison of tht! Arrhenius, Van't Hoff and
Thompson Te~rature Coefficienta
.Summary of Te;n~r ature Rate of Develop!llt:ln t
Exper iroon ts wi tl1 Salmonidae Egg a
Water Temp3ra ture, Incubation feriod and l-'er Cent
Mortality of Gret'n River Chinook Salmon Eggs
in Ex,Fer.iJnErlta I and II
Temperature Coefficients and Threshold Tempera-
tures for Chinook Sa.lmon from Green Ri'v·er
Reared at Constant TemJ>t!ratures (.2 lJB.gea)
Water Te:n~r ature, Incubation }l~riod and Per Cent
Mortality oi' the ChinooK Salmn h5~s of
Ex. fer i.tmn t II I
Observed and J1.djusted Tem~eratures Cllld Days to
50 f'er Cant Hatch for Chinook Salmon Eggs
Reared at Changing Temperaturee, ~rilMnt III
Estimated Thre ahold Ten,t..era. tures and Confidence
Limits for E.Jtl.'erimen ta I, II and III
T8Lilpera.ture Summation Constant, k
Water Tem,t..e rature, LlCubation Period and Per Cent
Hatch of E.lq.>eri.ment I Sublets
!-age
5
8
11
23
31
35
39
46
53-54
67
68
73
15
77
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
List of Tables
(continued)
Table
16.
17.
18.
19.
20.
21.
.a.
Water Te::lperature, Incubation Period and
Mortalit-y of Chinook Sal..mon Eggs .keared at
Constant Ta~~eratures at the Entiat
HatcherJ (Burrows' Data)
Abnormal Fry in Lots Reared at Constant Tempera-
tures
Average Weights and Lengths of Eggs arxi Fry
from Exre r:imEI'l. t I
Vertebral Counts of Green liiver Chinook Salmon
Reared at Constant 8fld City WAtP.r Temf'era ture
Vertebr~ Counts of ~acr~nto, SKagit, Green
and Entiat River Chinook Salmon of
Ex.perim.-n t III
Numbers of Uorsal Rays for Chinook S&J..mon in
~er i.ment s l, II and Ill
NUJ'llbers of An.E-..1 Ra.y s for Cninook Salmon of
Ext-erimtn ts l, II and III
Page
88
91
92
100
102
109
ll2
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
I. INTRODUCTION
To the fish living in the Columbia River and other Pacific Coast
stre&ms. ohanges are occurring :\.n the environment because of the impound-
ment of water for hydroelectric power, the increase in pollutioni and
the diversion of river water ~or irrigation. rn the future the use of
river water to cool nuclear reactors also may change the environment for
river inhabitants.
One of the environmental faotors th&t h ohanged is nter tanpera.-
ture, a factor to which fish, a poikilothermic anim&l. respond readily.
Usually the change is to warmer water and temperatures that are not favor-
able for the surTival of salmon<) An exception is the Shasta Dam on the
Sacramento River where the withdrawal of water from the storage reservoir
is from a level that is below the thermocline. AB a oonsequencea river
temperatures below the dam during late summer are now as much as 20"F
lower thAn formerly (Cope. 1949 and 1952) and salmon ant! trout produc-
tion has increased (Moffett, l949J Smith, 1950).
The changes that are occurring in the Columbia River and in other
Pacific Coast streams may subject salmcm eggs and young to unfavorable
water temperatures. a condition which also could ocour from the early or
late arrival of the parent fish upon the spa"Wlling ground, or from abnor-
mal wsather conditione. These ue some of the reaaons for accumulating;
more information on the influence of ttaperature l.lpon •lmon eggs and
young. Speoi.fioally, the objeotl vas of these experiments were to meas-
ure the etfeots of temperature upon chinook salmon, Oncor~chus tshawyt~
soh& (W&lbaum). in regard to:
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
2
(1) the rate or EIJlbryological develo~nt
( 2) mo.rtall ty
(3) occurrence of morphological abnormalities
( 4) grmrth rate
{6) the determination of the n\Ullber of vertebrae, dorsal fin rays,
and anal fin rays.
The period ot observation -.s tram the egg to the fingerling stagec»
Objeoti ve ( 6) above 'IF& a included for the purpoae of making e. con-
tribution to the limited inf'ormation now available on this subject for
the Paoifio aalmo12. and because of the current intere,st in the use of
meristic ohe.raoters as on• of the means of identifying the raoial orig.1.n
of salmon nO\¥ being oaught in the of'f-ahor• waters of the North Pa.' ~f'ioo
After Heinok:e• a in-n.tigationa in 1898 meristic oharaotera have been
uaed trequently to 14ctity racea ( ••• P• 96}. Heinoke identified groups
ot barring aa raoea when the ditf.-enoe in the oounta of vertebrae be-
tween groupe •• atatiatioally d.gnitioant.
Variation in the DWDbar ot vertebrae between indirldua.l fish is a
aouequnoe ot both genstioal and errvircmmmtal t'aotora. In field sam-
plea neither senotypioal nor phenotypical ~r1ation oan be determined
bee&UM the paat history ot the individual ia not known and therefore
the nrte'br&l oount ot the pu-ente IUld. the cmrlrcmmental factors intlu-
enoing Tertebral t~t1on are not knownc Bawenr, in laboratory
aperiaeDt1 auah ae tbeae. ICDe ea1d.ma'te ot genotyp1oal and phenotypical
ftriat1011 can be -.de aDd 'bhia 1Dt'01"JBB.t1on ia important to interpretation
ot reaulta ot l"&o1al atudiee baaed Clll tield ample a.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
The chinook salmon were seleoted for experiment~tion because they
are economically important, are rea.red extensively and are availableo
In the Columbia River and other rivers of the west ooast of the United
states that are most affected by d&ms, pollution and water diveraionso
this species is often the most important. OVer 30 million f'f"'j and
fingerlings were reared and relaat~ed by the washington State Department
of Fisheries in 1953. In addition, this speoiee is readily available
for experimental use, either from the Fiaheries center, university of
Was~..ington, or .f'rom nearby hatcher les of the washington state Depart-
ment of Fi8herieso
The chinook ea.lmon, also known as king, spt"ing. quinnat, tyee or
blaokmoutng is the largest of the Pacific Coast ~lmon and ueually
spawns in the large rivers. Spe.wninr; occurs as oe.rly as August in
Alaska and as late as December in Ce.liforniag the extremes of its gee-
graphical distribution. Tho egr.s are deposited in a nest in the £;ravel
of the stream bed that has been dug by the female, are fertilized by
the ma.le ~ and then a.re covered with grc:vel o While in the gravel, the
eg~:;s hatch. The young f1·y• work dO'I'f'llWard into the stream bed, a.s they
are neGatively phototropic. When most of the yolk has been used, the
fry B!r.erge from the gravel and soon gtart to seek the:ir own .food., The
seaward !rip;ration may b.,gi:'! .tm.-nediately. but l)ften 1 s delayed ur;til a
few months a:rte::-.f'eedinr;, and occasionally !U much as a j"'ar. T\~a :1hi-
nl)ok esbnon live in thtJ vea until maturity; whloh is usl.l.B.lly at an age
of 3-t! or 4-~ years, but rray vary f'rcrrn 2t to e?! year3. At maturity the
•"F'ry" is the stage which follon hatching and during whiah. the yoll: sao
1' ab1orboo; when feedine bet;i:'.g. the fry becano "t'ingor Ltngs,"
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
4
c;hinook salmon, weighing lu tt.:~ 50 pounds, returns to its na.ta.l stream to
8pe.wn and die (Clemens and Wilby, l946)o
Although the difference in spawning ti.me from Alaska to California.
may be as great as fo'Ur or five months, spawning ooo ur s when stream tem-
peratures are fa.lllng. Jordan and Everman (1896) write, " ••• it spawns
in August a.nd early September whm the water has raaohed a temperature
of about 540p-." TemPtJratw-ea observed at the 'time that chinook salmon
are spa1ming in some streams of British Columbia, washington, Oregon,
and Cali.forn.ie. are reoorded in Table 1. The water temperature during
incubation of the egg is descending, with minimum t«nperature occurring
shortly before or a.tter hatching, and ia risinG during the late fry e.nd
early feeding stages. The water t.mperature oyole in the hatchery at
the univez-sity of 'Mlahington as shown in Fig'Ure 1 is typioal ot the
anntal water pattern of stl'eU18 in whioh chinook salmon spawn in the
Puget Sound region., However, many streams in other areas of the Paoifio
Coast have minimum water temperatures of ~2° or 330f.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
5
TABLE 1
1e~peratures of Some Pacif1c Cuaet Rivera at the Tim~ of Spawning of the
Chinook ;;)almon
----·-·----~-
--------o, 1 eraperature, ,
River Wonth r.rriva.l .::if.i&wning l:ieference
Necha.i{o (B.C.) .::ie_~jt&11.ber 59-61 58-55 Hourston, 1953
~uesne.l (B.C.) late Aug. 60 50 Jackson, 1953
North Thompson (.b.C.) early Se.!Jt. 60 II II
Sc uth Thompson (B.C.) .nid Sept. 62 58-55 II "
Cc1umbia ( wVash.) (.)ct.-Nov. 62-50 ulson-roster,
.Skagit II September 52 See pCJ6e
Entiat II October 52 II II
u-reen II Cctober 50 II "
Toutle II October 58-42 Burner,
l • .il1&met te ~Ore.; Jet-tember 64-43 Matson,
Sacramento (Cal.) GctoLer 54-52 1-'elnar,
~Also includes a record of temperatures for other tributaries of the
Co1<.llllbia River
16
17
17
1951*
1948
1953
1956
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.~ I 7011 lr 65 a 60'" 55!-So a I 1r 4511'-1, k---~="''-··~ ,L~'-·· J ~·---~--J SEPr OCT NOV DEC JAN FEB MAR APRIL MAY JUNE JUU Figure 1. Average Water Temperature by Weeks at the University of Washington Hatchery, November 1950 to Ju~ 1953 0\ .J AUG
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
II. PLAN OF :&UER.I J.ENT S
The ef'feota (If temperature upon young ohinook salmon have been ob-
serv8d in three ..xperimenta in three suocesaive yea.rs-1951-62 .• 1952-5~.
and 1953-64. These exp•r1ments differed in tour prinoipe.l aapeo"t;sa
( 1) water temperature pattern. (2) aouroe of water, (3) raoial a took.
-.~d (4) number of pairs of parents (Table 2).
Two or the many possible temperature patterns were oonatderedJ tem-
paratures were ~ithor malntain~d at a ernstant lovel o.r ~re altered
throUfi:ho:rt tha experiment it1 a manner sL"r,ila.r to thn.t w~1ich occurs mxier
those to whi~h the ~ish a.r~ ex))osel! in oo.t·J::-e, for :.tis possible that
~lcrr-...nl developnent o~ egt:s a:rn f-:-._; is acl just~d to th.e natur&l 't'Jn.ter tem-
pers.t. 'Jl' e pat tern. T ~c nnture.l 1rn.t er ten?ern. ture s a.t the +; iJr.e of t-he
de~ositior. of eg;r;s in •he ::ravel !U"!l declininb and cor.ti!".1le to dodine
duri~ the inoubati on period.. Hate~ :1r: oocars about t!cs t 1m8 t:,at the
water tem.P6ra.tares re&c~ their lcwest level in t.he a.r:nual temperature
cycle, but emerger.oe of the try fran t~e ;ra.val do"s not occur until
;"R.ter ten;;oratures are ir.crea.s1nf HOW'evo:r. in a.r. C!:leperi.:nent in 'Which
the t(Jllpera.ture is :na.lntai !:ad at a consta~ le~el, the average tem-
perature may not be reh.t,d to the observed effect if t~e oharaoter being
o,served is influenced by t~?arature for only ?art of the observational
per1 oo. Beca.uee the oritioal periods for t~e eunook saL.,.,.on were !'!ot
known, t.emperattU"es of the bts for the first two e:otperkents were con-
s'ta..'ltJ for t.he third oxperi.r..ent temperatur~s were altered tr.roughout the
experiment in a manner sonewhat similar to the temperatures that ooour
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.TABlE 2 Summary of Factors .tielated to E.x}Jeriments I, II and m Experi-ment Date Stock I 11/15/51 -10/2/52 Green II 10/13/52 -l/l../53 " III 9/ll/53 -J/ll/54 Skagit 10/8/53 -5/8/54 Entiat lG/XJ/53 -5/?.'54 Green 1G/30/53 -5/8/54 Ss.cramento *Tap wat.er as race ived at th e hatchery **34,40,45,50,55,60,62~,65°F 145,50,55,57~,60,72~,65,67~0, ##45,50,55,60,65°F Water .SU}Iply City II Well II " " 0 Temperature Range, F No. of Temp. E.xver. Control s~. Pattern Lot a Lots* Pair a ----·--Conetant 34-65** 54-42-69 1 II 45-671 60-47 1 altered 45-651111 56-53 3 " II n 1 n II II l II II n 4 Ave. No. No. of of Egga hJ..er. in Each Lots wt 8 547 8 518 5 504 C) 5 559 5 452 5 586
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
9
under natural conditions ..
B:tperiment I
In the first experiment ~ght lots wore rMred at ~on~ant tempera-
tures in water f'rCIIl the city ua1n, and all eggs were taken fr~ one pair
or chinook: salmon of the Green River raoe. 'I' he experiment was 1n1 tiated
November 1~, 1951, and concluded Ootober 2, 1952?
A pair of mature ohinoo'k salmon from the Green River Hatchery of'
the •shington state Dftpnrtment of Fi sherine a tlrenty-f'ive miles southeast
of S"la.ttle, v.ra.c tra.nepartod to the uniVOJ'sity of •shin~on in a live-
tank a.nd than spawnedc The pRir used for spawninr, arrived durinr: the
lAst \'reek of the 1961 run of chinook salmon to the Gr~ Ri vt'lr Hatehory-
whloh is locate-d at Soos Creek. At this time the dni~/ tonperature of
Soos Creek varied from 42° to 460,. A.ft.er heinP". spawn,C,. tho pair vrere
photo~a?hed (Fi~? 2) ~d ~ize ~ensur~ments and counts of meriot1o ~har
acters were mad~ (Table 3).
The or,;: a from thi G !Jair werry d1 v1d~d into ei r-ht experimental a.nd
four control lots a.vera.r.inr, about 660 pM" 1 ot. ()le of the eight exr~r
in~ntal lots was plaoed in eaoh of the follo~ng oonc.tant temperatur~ss
~4 °1 40°, 46°, 50°, 55°. 60°, 8~ and 65<7.. A JNU:t.mUM tmnperature of
A5oy.. Yffl.s ohoa.,n bf!'oause in an earli~r t:rie.l cxperi:-1.ent e.ll e~P'8 at a.
oonlrtant tetn!1era.ture of 73"? and 67~ had di~d. 'T'he oontrol lots were
rearfld at tha water temporat.ure as l"*'oei vcd from the o1 ty Tlw.l.in. which
w~s 550p at the be~innin~ of the e~perin~nt and 41~. th~ low for the
Y"ar. 9.t time of hatching. Gre.phe of tho temperature history of the lots
of the first ~peri.ment are dl'lOWll in Figura 3 o
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
10
Fic;ure 2. ChincoK. ...JaJmon, 1 0renU; or' the Ex.f-eri.ment I Ec:;g Lots
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
11
TxBLE 3
lieasuremant Data of the farents of the Lots for
fuq:;er iments I and II
·,Ieight (a pawned), kg
Length (fork), c.m
Vertebrae
Dorsal rqs, all
Anal II II
Branchioatega1 ~ys
---~------------------
0ill rakers, first arch
Pyloric caeca
Scales, latere:U line
"
"
II
II
above " II
below " "
weight before water
hardening, kg
dia.llBter after water
hardening, em
Age (scales)
------------
hperiment I
14ale
3.83
95.2
67
1)
17
15
9 + 13
155
142
31
3+
Female
8.54
92.3
68
14
18
17
9 +12
139
29
7024
3cl8
0.93
4 +
--------
Ex~riment II
Male
B.f?/7
94.0
67
13
18
14
10 +15
134
3+
Female
9.14
95.2
66
lJ
18
16
9 +13
140
.30
6864
2.83
0.89
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
12
65~119
117
$5 -------= .. ·-· . . . . . .
.. 50-• \
l
i
E-t 45
35 -
. . . . ·-.....
' ..
\
' . . .
. . . . . . .. . . . . . . ·---. . .. . ...
.·
114
113
/}1
30 ~~--,-----~-1-.,•o.··•. ,_.-,, . .__L ___ ,,.--.-.... L--.---L......-·-----
0 5 ].0 15 20
Weeks After Start of ExperiDmit
Figure 3. Lot Teapera:tures far E%periJaent I
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
In thi8 experiment ei~ht sublots of tOO e~[S eaoh were renoved from
the controls to either & higher or low~r temp8ra:ture. After 3·} weeks e..t
oi ty water temperature thre6 of the B'lblots wera triUlsferred to oonste.nt
0 '0 water tempera.tures of 60 • 6q· anti 650f 0 Three othe!' control sublots
were trt.neferred a.fter 2 P 2~ and 3~ weeks at city water Um1pera.ture to a
oonstant t8lnperature of M ~. '.I'lro addi ti anal sub lots were held at 40°
and 46'1o' for four weeks e.nd were then moved to a oonstant temperature of
so"F.,
later, another seoti or. waB !tdded to the first experiment for the pur-
pose of observing the effects of tmperatures of 60'1o' a.nd hi~her upon
finrerlinr,s. At these temperatures none of the oricinal lots survived
to !'ewdirt(;. To ~stat>l1sh ~his part of the f:IJC.?eriment, the cor:trol lots
ware pooled on May 1. 1:152, and four r&ndom lrts of l.OO each were with-
drawn~ Of the!!le fo·,rr ~ on•3 waG let't at. th~ temperature of the city water
'.Lnd the others were trt.nsfnrrf3d, a1"ter g;rad·-19.l tampering. to constant
temperatures of 60°, 6 7° and 74 '1 .
~perimant II
The second axperiment duplioated the first by the uee of one pair
of' chinook salmon from th& Green Ht ver l"aoeo a.nd by the inoubati on of the-
egk;B in the se.m~ water a upply wd at oonsta.nt temp3ratures o
The aeoond experiment was ~tarted on October 13 wtth a pair from
the ear 1y pQ.l"t of the 1962 rw which waa late in arr.i ving at the Green
Rl ver Hatchery because of we.rm we&ther (Fallert, 1952) ~ The temperature
of the v.tater at the time the .fish were taken for spawning was 520p"
Size measurements and counts of meristic oh!U'a.oters of the two parents
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
14
for the 1962 experiment are given in Table 3 ~
Constant tmnperatures in the eight exparir..ental lots were 45°. 50°.
55°. 57~0 • 60°, 6~0 , 65° ~d 67~ .. These temperatures differ fran the
first experi~nt by the ~ssion of the 34° and 40° lots and the addition
of the 57~0 and 6~ lots .. As in 1951 t 1-:ere were fo~ control lots,
but sinoe Experiment II started a month earlier than Experiment r~ the
inoubation temperature of the controls was higher in 13529 The city
water teiP.perature at t~ ~sin:.ing of the experir.1ent was 61 '? and at
hatching was 55'7, the averar;e inoubat ion temper at .u-e for the oontrol
lots bei.Ilb 11 der;rees hi ;her t'ruul in l9~l, The ter.:pe.rat·..~r.., !-~s":'o!"·; of
the Experiment II lots is sht1'vn in FiQ;Ltre 4,
Exper:l.Jr.ont II "Na.S oor.oluded. unexpecterll:.t on Decemt)er 28 by &:n acci-
dent that oau5ed a. Great in(~rae.se in :nort.al ... ties in all lots. The cause
of death was a hJ .::;h 9:1 ~ tho water su?pl;t' c:-oo.ted by a chanr,e of char-
coal in the filter system~ The details c~ this ~ccident have been re-
ported by Seymour and Donaldson ( 1953). At the t1.-:1e of the accident the
lot at lowest tenpere.ture had ju3t eo-npleted ha. tohinr,, The exper1Dlent
'W.S o<mcluded at thls time, aa there was 100 per cent mortality in some
lot6 and injury of unknown extent to those that did surv1 ve ..
Expor iment I I I
The third experiment wa.a basioally different fran Experiments I and
II. The te~r.peraturee ware ohanging rather than constant 1 the water was
frrun a new aouroe. a well that had ba~n drilled to provide wat~r of lower
temperature for the hatchery; egr;s from four r&oes were ueedJ and the
eggs far one raoe were from one or more pairs of parents.. On Septtlllber 11~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.g.. .. I i-t ',~/\A ~ il ~ v V\ ..,1'1 'I 1f.u.J ~~ A,H I 60 j S$ f I 1 sorr... ~ -~ 4S i '\ \ \ \_-_,"' /~, '---16 \......, _____ -.... ...~'"'\..__./" J,-14 it'l ~~L·---~=--=--=--_h_-L __ ......::::.:o _l_---=--........-.....!---....J. """-~--J....... ____ ,_-J.....---L--.. ~-----.__-__ 1 ----=--.......J...._ ~ 2 7 12 17 22 27 32 37 42 47 $2 $7 62 67 72 77 82 87 91 DAYS AFrER START OF EXPERIMENI' Figure h. Observed Tempers.tures by Days tor Lots of Experiment II ~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
16
195~# the exp~rimant was started and was continued untll May 8, 1954~
For the purpose of evaluating racial difference• oaused by tenpera-
ture. esp&eially the rate of development and meristic characters. eggs
were obtained !'rom four rivm-ss the Skagit, in northwestern ?alshington1
the Entiat, a tributary to the Columbia River in e~stern Vlllshington; the
Sac~ento, in california; and the Green. near Seattle (from which eggs
were obtained for the two earlier axperiments). Unsuccessful efforts
were made to obtain eggs from Alaska, first frcu11 the Naknek River, Bri s-
tol Bay, &nd later from Alex..nder Cr•ek near Anohora.ge. It was considered
desirable to use Bristol Ba.y fish a.s representative of the races of chi-
nook sa.lmon near the northern limit of their distribution. Because the
eus from Alaska were not available, ages from the early Skagit River
run were obtained.
The eggs from the Sk:agi t River chinook salmon were taken on Septem-
ber 11.. The part.mtsJ three females a.nd two males. were gaffed tram the
Marblemount spawning E;Tounds. The water temperature at time of egg-taking
was 52~. After fertil1tation and water-hardeni~g the eggs were combined
into one lot, placed in a large thermos j 1.1g &nd taken to Seattle e Water
temperature in the jur; upon a.rri val in Seattle four hours later was 59~.
One-half of the ee;gs were then forwarded to Mr. Burrows at. the U. S-Fish
and Wildlife Fish Cultural Laboratory at Entiat, wasnington. and the re-
maining half' divided equally into five experimental and two oontrol lots
of about 500 eggs each.
The eggs from Entiat R1vor were obtained on Ootober 6 fran & single
ohinook female about 15 pounds in weight. After tertili£ation the eggs
were W&.ter-ha:rdened for three houri and were then plaoed in the BaJne
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
17
~ype of the!"TI\os ju_e: B.R had been u.ettd for the Skag;i t Ri ftr eggs. The
Entiat Ri1fer nter temp81"&t1.tl"e ,...., 520po at th'l..s time:-The jug was pre-
oooled with ~rushed ios IUld. &.rter bein(S till~ with er,~s &.nd water, was
oovered 1f'ith 1nsula.tlne; aterU.l. Upon arrival of' the jug J.n ~eattle
four hours lAter. the ft~r tan.perature 1na1de was 49'T ~ The eggs were
di Tided eque.lly 1nto six late or about 660 e&oh.
The eggs of the ohinook salmon from the Sacramento and Green Rivers
were taken Co'tt'b~!" ~0 ~ T hA Saorlllllfmto eggs were spawned in the morning
at the Coleman Station &nd flown to Seattle the a&me day. One-half of
these egc;a was ta.ken to the Fish Cultural Laboratory at Entiat., The
eg~:;s were shipped frOlll California in a special oontainer w1 tt• ice. and
upon arrival in Seattle the temperature about tho eggs was 3Bor. On the
de.y ot' spawniug the temperature of the sacramento River at the pla.oe
from which the salmon were taken Vll.ried fram 52° to 54~. Four pairs
of chinook sa.lmon were used and the eggs were mixed before division int.:>
experimental a.nd oontrol lots of ~bout 680 e&oh. The fish in the Sacra-
mento River on October 30 were the first of the fall run of ohinook
salmon a the pee.k of the see. son's run was expeot.d later.
The Green River egr,s wer6 taken fram one female and fertilized by
one rns.lee 1'he 6G£:S were transported to the University in a thermos jug
in whioh they were retained ll'ltil the Sacramento eggs arrived, The tem-
perature in the jug increased during this period from 52° to 59'1<', Both
groups of eggs woere h&ndled in the same manner after being removed from
the shipping contt..iners-The average number of Green River eggs per .iot
wa.e about 460o
The temperature pattern ohoaen was that which olosely reasmbled the
pattern expected in nature. The average temperature of the oity water
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
18
in the hatchery of the Sohool of Fisheries from November 1950 to July
195~ was the pattern (Fig, 1), but the actual temperatures were at a
lower level for some lotso
0 For the .five experimental lots the starting temperatures were 45 ,
50c; • 55°, ao 0 and 660-p-. After the start o.f the experiment the water was
cooled one degree fi'Very five da.ys to a temperature of 34'7, The egrs
were held at that temperature for twenty daJ'S, after whic!: the t€1n.pera-
ture was increased at the rata of one degree every five days-The
temperatu:t'e history of the Experiment III lots is shown in Figure 5.
The water source for the third experiment was from a well, whereas
oity water was used in EXPeriments I and II, This was of importance in
~ respects, First. the tsmperature of the control lot which was reared
in the tap water was pr~ctioally constant, with a range of only two de-
grees, from 56° to 64~. secondly, this water a&rried a hir,h organic
load which resulted in the very ra.pic auoumulation of slime molds, algae,
&nd prototoans on the egre, in the tanks ~1d troughs, and in the coils
of the cooling system. This necessitated more handling of the egLs than
was desirable &nd interfered with the flow of water in the refrigerated
tank: so
ay using ultraviolet light an effort was made to oontrol the slime
molds, algae and other organisms growing on the bottom and sides of the
troughs e.nd tanks. Three ultraviolet lights were placed across the head
a£ a. trough without fish, a few inches above the water. The water depth
was four inches. After three weeks the mass of organisms beneath the
l&mps was only slightly leas than in other parts of the trough& there-
fore the use of' the ultraviolet light was discontinued.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.r}&4 ... I ... 65 ... , ' ' ' ' '• ' r.o~S 60" ' ' . ' ' ' ' . ' ' . ' ' ' ' Lo~ ' ' 55~ ' ' . ' ' ' ' . ' ' 'o. ' ' ' " ' " ' ' ' . Lot',J ' ' / . ' 50" ' ' " ' ' ' . ' / ' ' ""' ', . ' ' ' ' / ' " . ' ' ' Lo'2 " . ' / 45' ~ ' / ' . "' ' / " . ' " ' ' / / . " ' / / / . / 40-"' " ' / "" / / / / "' / / / 35 ~ "' " ,, '/ o hatching -_J'"----' ~=-----------=-,-_;---·! ·-0 25 j 50 I 75 '· j 125 I 150 100 NUMBim OF DAYS AFI'ffi START OF EXPERIMENI' Figure 5. Predicted Temperatures from the Beginning of the Experiment to the Fingerling Stage for the Lots of Experiment III 175 / / 1;0 / / .-L 200
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
20
Mortality in all lots of ExperL~ent II! in the late fry and finger-
ling stages was unusually high, and although the speoifio reason was not
found, the oauae was believsd to be either directly or indirectly asso-
ciated with the well water o }fortali ties were especially high in the
Green River egga a.nd. moreover, they oocurred earlier than in the other
lots, indicating that other factors aeaociat8d with the condition of the
eggs themselves were contributing to the mortalities.
Two lots, Sk 6 and E 6, which were reared inadvertently under abnor-
mal conditions are not included in the discussion of results obtained
from the other lots. At the beginning of the experiment these two lots
were in troughs containing well water in the main hatchery and were not
transferred to the controlled-temperature hatchery until November 14.
On October 29 the dissolved oxygen in the water flowinc into the trough
with Lots Sk 8 and E 6 was found to be only 3.0 ppm. This condition was
oorreoted the following day. However~ by that time Lot Sk 6 had been in
the low-oxygen water for 48 days and Lot E 6 for 21 days.
The si~if1oanoe of low oxygen tension was not appreciated until
the data were analyzed some time after the oanolua1on of the experimento
Then it was found that in the two lots affect~. the incubation period
wae 18 per cent longer, the increase in the average number of vertebrae
was ae muoh ae 2o6. and the increase in abnormal vertebrae was about 10
p.r oent.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
III o MATERIALS
The hatchery and equipment of the School of Fi !lheries in Roam 1.24 ~
Fiaheriee Center, -...roe used for these experinents (Fig. 6). The roan is
provided with six tanks and four troughs with tap water and heated tap
water supplied to eaoh0 Refrigerated t&p water is supplied to four of
the tanks.
The tanka are made of ba.ked enamel with a. Thermo-pane glass front
L~d are 60 inohas long, 2~ inohes high and 12 inches deep. Protection
from temperature oha.nge war.~ provided by two inohee of cork on the ba.ok.~~
bottom and sides, an ins·.<lated lid on top, and the 'I'hermo-p&ne front.
The oonor~te troughs are 15 fest long~' 1.2 inches Wide and 8 inches deep.
water. For the first two experiments the supply water was frcm the
city maine To remove ohlorine gas tha.t is a.ddt"d oooasiona.lly to the city
water. all water for hatchery use wae ptUaed through a oha.rooa1 filter..,
In Experiment III well wa.ter replaced oity water. The temperature
of the well water W&l 68° to 56ot, a favorable temper&ture for chinook
salmon. Analyses of both the oity and well water is given in Table 4 ..
For the low-temperature tanks the tap water was oooled in a Temp-
rite Instantaneoua Cooler and temperatures down to freezing were avail-
able, Heated water was provided by running water through a coil in a
stoam jaoket. Intermediate temperatures were obtained by mixing warm
and oool water, using a. shower type valve, either Powers model 34504
size C-20 or Powers model HVE.
Tra.ye. A tray was designed so that the egr,s could be kept in order
and examined individually, if necessary, with & minimum of handling~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
22
Figure 6. The Controlled-temperature Hatchery Room,
Fisheries Center, University uf Viashin15ton
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Date
23
'l''l<L::-4 ll.l.. -
Mineral Analysi:5 cf City a.nd Well V.at-e:r
City
Wi:!tt::r;,
l/7 I s.2
City
';~a.t~rl<
1/5/5)
City
Water o:-'~"
l/C7/54
iiell
Wa.ter~:-;:
l/29/5L
h::.Ft:3 _Ler 'Hll ic n_· ______ _
Total .::>olids
Silica ( Si02)
Iron (Fe)
.11.iu.ninum ( Al)
C a.l c iu:n ( C a)
.Magnesium (Mg)
l-otassiwn ( K)
jc•:iiwn (Na)
bicarbona.Le (HC03)
Sulph;;.te ( 30 4 )
Chloride (Cl)
'futal Hardness ' \as Caco 3 )
i~.lkalinity to rhenolfJhtha lBin
It It :Jetnyl Orange
pH
42.
) ' .:. .(
.. ) . :::
6.8
\J.'l
1.9
19.8
G.
2G.
* Analysis by Seattle Water Department
u.
ll.
,,,04
,J.Ul
~.8
l.
C,J
1.88
2.9
1.06
2(;.1
G.
24.
7.4
** 11 11 A}JIJlied Fish::r ies L<s.boratory
))7.
'7 ' . lL.
2. 15.
t;u. 117.
..::J. 1r• Ct' •
7.6 '!.4
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
24
Tha tray oonsistf.td of glass r.::d.s st:.,t:;a.li" t~·= ln !1. 10-by 12-inoh wooden
fr&me. The rode were spaoed 80 that the egr-s were supported.~~ but the
larvae" upon hatohinf, dropped through. One tray held about 600 chinook
aa~on egr,sg 20 rows of 30 eggs each.
Da.mboards. Sinoe in the third e.x.perimer.t there were as rrany as
four different lots in the same te.nlc,. it was neoess&l7 to divide tr1e
tank into oompartm8nts and to have th~ ds:nboard between oompe.rtments
impassable to the fry and finEerlings. The oampartments were made tight
by wed~ing a damboard against a half-in~h sponge rubber r,asket on the
sides and botttml of the t:..anlc.J The da.:nboe.rd was made of two plates. one
inoh ap&rtg which were perforated at the top on the upstream side and at
the bottam o~ the downstre~ side. Suooessive d&mhoards differed in
hejght by one-half an inoh, with the higher damboard nearer to the head
o.f ths tank. Building and s.rranging the d&m.boards in this manner oreated
oiroulation of water throughout eaoh compartment.
Therm?gra.phso Tank and trough temper&.tures were constantly reoorde l
on ~ oirouh.r ohart, a thermogram. by means of a seven-day Bristol Re-
oorder. Temperatures w~ra oheoked daily with a o&librat8d mercury ther-
0 mameter whioh oould be read to 0) 1 F ~ and a.n adjustment was made to the
recording pen if the pen was in error~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
IVa MEtHODS
Egg-taking. In Exp$riment I the egr:s were taken by incision and
were fertili&ed in a pan without water, After fertilization the eggs
wer"' divided into equal lots and plaoed in pans in the tank in whloh
they were •.o be reared~ During the next two hours the eggs were ;n.ter-
h&rdened and temper~ with the temperature gradually changing from that
of the eggs at time of fertilization to that of the rearing temperature,.
the greatest change being 14 degrees F&hrenheit. The eggs were then
placed on the glass rod trays. In EXperiments II and III the eggs were
wa.t&--hardened fer two hau:rs at the site of egg-taking and then were
transported to the Sohool of Fisheries. Illmlle<iiately after arrival at
the School thtY.f were divided into lots and tempered for two to four hours
to the rearing temperatures ..
Handlin§• Egg• and fry were moved only when absolutely necessary ..
The tanks ware cleaned with a siphon and the dead egg a and young were
removed without disturbing the remaining eggs or fry. Except for the
removal of the sight sublota in Experiment I, the egLs and fry of Exper-
iments 1 and II were undisturbed. A.s mentioned above, this was not true
in Experiment III. Accumulation of organic debris and mud made it neces-
sary to agitate the eggs gently about once a week to prevent smotheringQ
Also. some egg lots were transf'erred fr001 one tank to another during the
eyed egg stageo
Mortality records, )dortali ty was oa.loulated from the number of eggs
and yotmg ramoved eJ'ter the first day. The removals on the first day
were mostly infertile eggso Dead eggs nre removed daily. but since
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
26
death of a.n egg is not neoessarily :.~.:M:~u1£<tely ap~ent~ mortality V'!il.l-
ues for any one day during the .acg stage may actually be greater than
shown in Tables 9 and 11. This souroe of error is not present after
hatohing.
The cumulative mortality was oaloulated by weeks for eaoh lot. The
losses were separated into two eateEor1ess one oategory was natural mor-
tality. the other ramova~~ aooidental deaths. The cumulative mort&l-
ity took into aoco·mt losses f'rctn both categories and W&s ocnputed for
n.oh ~even-day period. The oumulati ve morta.li t:: for week n was the sum
of three i terr.s, ( 1 L the oum1llati ve morte.li ty for week !:.:.!,) ( 2). the
sum of the dai ty natural mortal! ties for week ~J and ( :5) ~ the natural
mortality that would have ooo\.U"'red in week.!: among tr.ose removed for sam-
ples or killed aocide~tallyo It~ (~) was oaloulated by multiplying the
1'18.tural mortality rate f.'or week:!:, by the sum oi.' the number of individuals
that ~ere rsmoved for s~~ples or had died accidentally during week n and
the number of individuals that would have survived to the end of week
~· having previously been removed for sarnplas or having died aociden-
tally.
The per oent cumulative mortality was equal to 100 times the oumu-
lativs mortality divided by the number of eegs at the start of the
experiment.
Mortality for all stages WILe caloulated in this manner. Mortality
to the 50 per cent hatching stage did not include fry mort.alities tor
tho~e Wedks in wiuoh there w~re both egr, and frJ mortalitieso
Estilna.tion of the time to hatohlngo Ha.tchi:lg was defined as the
time when both the hee.d and tail w~e fl-ee of the shella At both high
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
27
and lew tsmperaturas only the heada of many larvae would break through
the shell and the larvae eventually would dieo suoh larvae were oonsid·
ered egg mortalities and were not counted a.s being hatohed.
The time to ha.tohing wa.a arbi tre.rily seleated as the time when 50
per cent of the eggs had hatched--the median of the hatching period-~
and could not be determined until completion of hatching. Other choices
would have been the time to the first hatoh, the last hatch, the mean of
the hatching period, or the mode of the he.tciing period_, At normal tem-
peratures the ohoic~ would have made little difference, since the hatch-
ing ?eri od 1 a ahort, At law temperatures the hatching period of some
lots was more than one month, with a peri)d of no hatching interveninG
between the extreme valuesJ therefore the time to hatching of either the
first or the last egg wae considered an 1nap~ropriate estimate of the
time to hatohinc.
The hatoh of ebge waa counted daily at ll a ..m. F'ram the daily
hatching records the time to 50 per oent hatch wae calculated to the
nearest 0.1 of a day, the last significant figure being determined by
linear interpolation. By this method of estimating time of ha.tohing the
error is relatively greater in those lots with the shortest hatching
time, i.e., those at highest temperatures.
Estimations of average temperature. The temperatures could not be
oontrolled perreotly and therefore it was necessary to measure the tem-
perature fluctuations in order to have a reliabl~ estimate of the average
temperature for the period of observation., The tempt,ratures were record-
ed on thermograma from whioh the aver~ge temperatures were calculated~
Sinoe the thermometers attached to the urlxing valves were o~librated in
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
za
F!.hrenhei t un1 t s, the thennogrltJils were c.ali bra ted in the same units.
The averabe tsmperature was calculated with the aoouracy allowable
under the conditione of the expertmento The reliability of the thermo-
meter • the accuracy of the thennor,rams. the tem~rature changes at vari oua
posi tiona on the egg tray. am the methods of oaloulation all influence
the aooure..oy of the estimates of &.vare.ga tem;>eratures ·J The thenaometer
u1ed to make the daily temperature readings and to correct the thermo-
g:rams was aooura.te to O.lOfJ tha thermo&J"&mll oould ~ read to the nearest
o.2or, anrl the r,reatest temperature difference between various positior.s
on the egr, tray was Oa20f.. A graphical method and e.n ari thmetioal method
were used to estimate average tem~ratures, but the l!l"ror in either
method was not measured" However, the tota.l error from all sources in
the calculated average temperatures is believed to be leaa than 0.60p.
Esttmates of average temperatures derived by means of oaloulating
areas under the thermogram or as derived uithmetioally from daily aver-
ages over short intervals are more accurate than averages fr001 daily
ma.xirnu.m and minimum temperatures. For Experiment 1 uverage temperatures
were estimated by computing the area enclosed by the thermoe;ram. From
the area. whioh was computed with a planimeter" the average temperature
for ea.oh seven~ay period was read directly from a. table~ In the table
the areas enoloeed by oirclea made by vari oua oonstant temperatures are
listed. For constant or nearly oonsta.nt temperatures the plan1..meter val-
ues are approximately oorreot. but for fluctuating temperatures there is
a.n error because t~ areas of the cirolea do not ohange linearly with
temperature a l'he planimeter method was chosen for the first year's ex-
periment. as the temperatures ware nearly oonst.aHt, espeo1a.li.y during
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
incubation.
For Experiment IIl daily avera.ge t.empera.tures -.are determined e.rith-
matioally from temperatureii dur-ing short intervals in which there was
81 ther no change of temperatura or a constant oha.nge in temperature. To
dl!'te:rm.ine the daily average, temperaturee during each interval. weie;hted
by the number of hours of eaoh interval, were averaged1 For example, if
the temperature during the first twelve hours of the d~y was constant at
50~, then steadily rose during the next three and & halt hours to 560f.
then declined steadily to 520? during the following two hours. after
whlch the temperature remained oonsta.nt for the rest of the day .. the
daily average temperature was oa.loula.ted to be
(12x50) • (3o5x5~) + (2x54) + (6.5x52) ~ -M -or 51 A -r'.
To exami M the di.fferenoe in e.V8T'8.~., tsnperature~; tfu\t would ocr 11r
from the ~o m~~hods deacrihed, ei0ht thermo~rams were e~leoted i~ ~~x
i:1. two of whiolc it wr..s 0xtrer..o ( irrer,:.~la.•· ":.~or:n.obra.m). Temperatures oal-
culated from the plw:i'Tlet&r me'\s-.u-aments avtJr.1.f:f3C G ,:i y-Jr oer.t les~; than
t"l'T.j:>orat~res o~lO'llated a.rit~atioally SJ.nd tfl.e error waB t.he ::;Q.m~ fo:--
atll!"ea th.ut were reoorded in l<:Xilerimants I and II~ Tha varianoe .,..,_,~
probably not r;rea.t 11 as teunpl'fratures were 1.A.sue.lly mainta.ined wJ thin one
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
30
degree Fahrenheit of the desired temperature exoept on oooasions when
the refrigerated water supply was reduced or out off. This ooourred when
ice or organio material aooumulatod in the ooils of the cooling unit and
when there was a power failure to the refrigeration unit~ The shutting
off of the refrigerated water was mowt frequent during Experiment III
and occurred four times during the experiment to the tank reoeiving 34'7
watero An estimate of the varianoe of the daily average temperatures
oould be made, but this would not indicate temperature shook~ as usually
the deviations fram tho tsmperature pattern were of only a few hours'
duration and changed the daily average relatively little. Although the
temperature shook e.ffeot t'rom daily temperature changes is not known. a
comparison of the daily llJI?..Ximum-mlnimum temperatures of the experimen+..a.l
lots with ma.ximum..-n.inimum temperatures in the Grt3en River (Ellis. 1953)
and the Sacramento Hiver (wallioh, 1901) shows that the experimental lots
were not exposed to any greater daily temperat·J.re changes than occur in
nature (Table 5)e
Preservation. Before being preserved. most specimens were placed
in urethane. They were then measured. "'eighed and ra.dio.:;raphed. The
preserving fluid was 4 per oent formaldehyde with 0.7 per oent NaCl.
Radiogra.phso Counts of vertebrae and fin rays were made from radio-
gra.phs e.Jld from stained specimens. All the fry and fingerlings that were
seleoted for coWltine; were radiographed, but the radiographs were not
readable for some of the fry with skeletons that had not yet ossified.
These try wwre then stained and the vertebrae and fin rays in many could
be ootmted ..
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
31 TABlE 5
A Comparison of the Daily Temperature RBI1be for the Green ane1 ;;iacr<Llento
Rivers and for Experiments Il and III
Temp. RB.nge, °F
0.5
1.5
2.5
3.5
4.5
5.5
6.5
?.5
8.5
9.5
10.5
11.5
12.5
13.5
14.5
15.5
16.5
17.5
18.5
i9.5
2C.5
Green
Oct. 195:2-
.M.:.y 195 j
Sacramento
~e}'t. 189·S-
F'eb. 1899*
Number of Days
2 1
11 2
68 12
30 27
55 14
18 14
35 10
13 1
8 1
" "-
1
2
1
3
1
1
2
* Only fuur observations Nev. 17 -Feb. 18
E..x.~-. II
Cct. 1952-
Jc..n. 1953
ExfJ. III
Sept. 1953-
.L!ey 1954
Number of T arlic Days
b6l 902.
120 220
19 72
10 40
3 19
1 18
1 5
5
7
2
2
2
3
2
1
1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
32
For radiographs of small fish soft radiation of high intensity is
ne~ded. Sharper and olearer radiographs oan be obtained from an X-ray
d1ffraotion unit than from a diagnostio type o!' tube because the betUn of
X-rays 1a snaller and there is less secondary radiation from the window
(Bonham and Bayliff • 195~) • A we.ter-oooled M.aohlett o-2 X-ray Diffrac-
tion Unit with a oopper targ3t and a beryllium window was uaed in these
experiments o
Kodalith ortho Type 2 film produoed good results andQ being insen-
sitive to red light. was conver~ient to use. A sheet of film placed in a
black. light-proof envelope was positioned beneath the X-ray tube s.nd the
fish arrt.nged on a sheet of cellophane resting ~n top of the film enV!9-
lope. The fish were blotted dry and covered with a. sheet ot cellophane
to reduoe further dry1.nr, whi oh sanet.imes resu.i ted in mo'V'Eilnent of' the fish
during exposure.
}. typical exposure for 30 two-inoh fish on a &he~ of 5-by '1-inoh
film with the window 24 inche& tra.'ll. the f'ilm and the u;U.t operating at
50 P[V (peak kilovolts), 12 l4A. (milli8.mperee) anc full wave rectification
was 4-~ minutes. The film was developed for 40-50 second a in Delctol di-
luted with two parts of water. For ls:Eer fish ( Hve-inoh) Type M X-ra.y
film WM.a used in order to s:1orten exposure time. W1th tha tube a.t 32
inohes e.nd operating at 50 PKV and 12 MA-exposure time was 25 seconds
and developmhnt time 4 minutes.
Satisfactory ra.diogra.pha have been mad& o!' chinook salmon rry as
amall as ~€ mm (fork length). These fish were reared at 40Vr tor 10
months and did not incre.se in length after haton1ng, whereae younger
38 nun fry in the yolk-sao sta.~e did not give Ga. t: e1'a.c.tory radiographs.,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
33
ossification appeared to be a fUnction of both size and age. Fingerlinea
were used for radiographs when availableo For those fry that did not
give a readable radiograph the staining technique wa.s also tried~
Staining" The technique used was that of Hollister ( 1934) for
clearing and dyeing of fish for bone study, with the modifications of
Tining ( 1944). Fry preserved in 4 per oent formaldehyde were washed in
water. bleached with hydrogen peroxide, washed. plaoed in 2 per oent
potae•ium hydroxide and then st&ined with alizarin (alizarin sodium sul-
phonate). Bleaohing was accelerated by exposure of the fish in the
hydrogen peroxide solution to ultraviolet light. From the stain the fry
were transferred to glycerin !'or clearing. , .. ertebrae and fin rays of
the glyoerin~preserved specimens were oour.ted with a Qissecting micro-
scope at a magnification of 7 x.
The ossified structures stained very di stir>otly. Of the two methods
the staining method was more effective for young fry, although all the
possibilities for the radio~raphio methodi suoh as voltage and amperage
changes and types of photor.raphic pe.per. were not explored. Aooura.te
oounts were easily made of the specimens of fingerling size prepared by
either method. but the radi O[;raphio mot hod n.s pre!' erred because the
radiographs provided an orderly, permanent record that was available for
rechecking and because the method was faaterc
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
V. EliVIROJDmfi' AL CONDIT I O!iS
The ~xperiment waa desi&ned to measure the etreot ot one variablell
temperature. other factors were assumed to be eithGr of no effect or of
equal erfeot in all lots.
OEygen. Analysie wa.s done by the b&do Winkler Method as outlined
by Ellis. Westfall and Ellis (1948).
On five occasions--January 10. JLnuary 25. May 26, October 1. and
Deoember 29 1 1952--axygen determinations 'J<rtu•o illli.de of the oi ty water in
all ten tan.ics and troughs with 8Ulples taken fram the int&.ke. the outlet.
the surface and the bottom. In all samples the dissolved oxyeen was
greater than 70 per eent of the saturation level &nd in most cases greater
than 90 per oent. There were no sigllifioant ditferenofJS in o:xygen val-
ues of samples from the intake or the outlet, the s\U'fa.ce or tile bottom
(Table 6)o
___ _,...--:i:~c> ...-~-n wa.ter uaed in Experiment lii w.aa pra.otioa.lly devoid of oxy-
gen &s it entered the reservoir tank at the Fisheries Ce~ter. the value
for cissolved ~gen in parts per ~illion being o~zo or approximately 2
per oent o1' Ba.turation. Arter spilling into the reee~oir tank through
wire mesh soreens. the oxygen increased to 3.4 ppm.
In the controlled-temperature hatchery the wa.ter was jetted into
the ta.nlcs and troughs, which f'urther increased the f'ree oxygen in the
water. Values ranged f'rom 7 .4 to 10.9 ppm and the per cent saturation,
!'rom 68 to 80. However, two lots in the main hatonery were in water of
low oxygen content, 3 ppm 11 on October 28. The tap water to the troughs
holding these two lots entered fran the bottom without mixing with the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.TABLE 6 Oxygen Content of Water In Tanks i.l. no Troughs City Water Well Water Tank or 1/10/52 1/25/52 5/26/52 lli/l/52 12/'2!-J/52 lu/2c/53 ll/13/53 Trough ppm %Sat ppm ~Sut !Jpm %Sat .fJPiil ~Sat ppm ~Sat .fJF.:l %Sat .tolJ!ll %Sat -~----------~----------------·---~---~---------1, inlet ~utlet l~L39 94 12.31 89 11.28 8~ 10.90 dG 11.20 7Cf 2, inlet outlet lj .02 99 lC,. 88 83 11.39 8') 8.65 68 3, inlet outlet 12."+9 103 h.-.59 87 lu.l5 cj 9.71 92 8.}5 77 4, inlet 1..-' 4t UO ~ outlet 12.6~ lC~ 1:~.30 99 9.31 96 8.49 89 8.30 75 5, inle t ll. 45 103 ou tl0 t 1~. u6 106 11.37 102 h.. 43 92 s. d. 72 8. OG '15 6' inl~t 11.59 107 wtlet 11.67 llO ll.vt lud lL.J7 95 8.34 2j ?.45 75 7, inld lC, .4::: 105 lC. bU 105 outlet 1G.27 103 ll'.57 104 9.73 98 6.52 68 2 inl6t 12.71 102 12.55 lul , --aut let L2.. 58 lll 12.50 l~H.J '). 2.l. '-11 '_-'. J 5 9c L. 7? (/1 7. 85 93 9, inlr1.::t <1.48 10J ') .. '5 97 7.90 b4 7.75 7'1 out et 10 i:,:.:.: 11.29 116 'vutlct 11.17 115 ~~.92 103 r;.,l>3 90 7.55 78 fl j, rnain hc.tchery 2.95 J..7 9.2.0 87
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
36
air. To coneot this condition overhead jete were installed.
Dissolved oxygen levels of 3 ppm or lower are hazardous or lethal
to fi:~h in lakes or streams e.nd 6 ppm or more should be present for favor-
able conditions (Ellil! ~ al., 1948). Low oxygen le'ftll also influence
the development of fiih eggs, as Johanaen and Krobh (1914) have shown
that levels below 60 per cent saturation delay the development of plaice
eggs.
Because the well water used during the third experiment was rich in
organio material, the bio~ogioal oxygen demand (BOD) of the water also
was determined. Three 250-ml samples or well water incubated five da:p
in a hatchery trough at the well water temperature of 66e>r had net oxygen
losses of o.s, 0.6 and 0.7 ppmo A fourth ~ple treated in a similar
manner, exoept that the bottle was deliberately loaded with organic ma-
terial gr<JWing on the bottan of the trough, had no tree oxygen after five
dayso Although there waa a positive BOD. the decrease in oxygen from.
inlet to outlet was no grea.ter than the measurement error, 0.2 ppm or
leas.
lfate:l" flow 8 Water flows were determined empirically for each tank
and aTer&ged 1.2 gallons per minute with a range of i to li gallons per
minute. The flow to the oold water tanks was liDd ted by the oapaoi ty of
the refrigeration unit but satisfaoto~ oxygen levels were maintained as
1ndioa.ted ~bow •
.@• The hydrogen ion oonoentra.tion was determined with a Beokins.n
Model H2 Glaas Electrode pH Meter. Values were in the range of 7.6 to
7.8 exoept on one occasion when the charcoal in the filter system was
replaced. This resulted in a great increase in the pH value of the water,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
37
a great mortality and the conclusion of Ex?eriment II as desoribed on
page 14. Aoidity in exo6SS of pH 4.1 or alkali~ity grester th&n pH 9e5
are immediately lethal to brook trout (Creaser, 1930).
Ught. Tha hatchery room for the tem.per&.Llre experiments is a.n
inside windowloss roam. All lots were exposed equally to the fluores-
cent lishts in the rooro~ During incubation s.nd throughout the try stage
the lights wert~ en about two hours a day durin~ the daily routine of
t.aking temperatures, removinr; dead eggs and young, ~· In the feeding
stage lights wore on about 10 hours eaoh day. McHugh (1954&) has shown
some evidenoe that visible lit;ht durinr, embryonic development of the
t;runio~. Lauresthes tenuis, inflc.umoes the number of vertebrae. Mea.n
vertebral number is relatively low in bright light» intermediate in sub-
dued lir,ht, and hich in darkness. In nature snlmon eggs are in darkness,
boinf buried in the Lravel of the stream bottom.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
VI • DISCUSSION AND RESULTS
Rate of Embryologioal Developa!nt
General expression•• (a) Hi•torioal. The rate of development of
poiid.lothermio &ni.TU&le variea directly with temperat•ue. An expression
or the relationship between temperature and rate or de-velopment has been
sought by many in the hope of olaesli'ying physiological prooesses a.ooord-
ing to the size or their ooeffioients. It wab hoped that the site of
the ooe!'fioients ·would :reveal, by oomps.rison, the chemioe.l or ph.ysi.oa.l
processes whioh ue th" foundation of the biolo~loal reactionth
Mathematioal expressions proposed to de<Jori.be the relati(\nl!hip or
temperature to speed of developm6nt may be classified aa either theoret-
ical or empirical. In the first category are van' t rrorr' s 'ho• Arrhenius'
l'• and Thompson's x or Q1 , all of which are~ ba.s:4..oally the aame equation.
The three ~uatione are compared in Table 7 &nd are shown to be of like
farm after logarithmic transformation.
In the Arrhenius equation temperature is expressed in absolute units
and in reciprocal form, but Belehrftdek (1935) pointed out that the recip-
rocal of the absolute temperature 11 practically a linear function or
temperature on the centigrade scale between the limits of 0° and 40°C.
Therefore, the expressions of Arrhenius and van't Hoff are virtually
equiT&lentJ both imply that a proportional increase in speed of develop-
ment produced by a given di!'ferenoe in temperature is constant throughout
the temperature range at which an animal may deTelopo If ll fits any par-
ticular eet of empirical data, Qlo should fit equally well and vice ~~
(Andrnartha and Birch, 1954 )a what is true of Q10 11 a lao true of x or Q1 •
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.Coefficient TeiiiJ6rature symbol Temperature Wlits Temperature difference Rate Symbol Ori6inaJ. equation Transformed equation '!'ABLE 7 A Comparison of the Arrhenius, Van't Hoff and Thompson Temperature Coefficients ----Arrhen:iua J1 X Kelvin x.l -x,.2 y . 1 l ) u(---Y 2 . e z .x.l x.2 yl 4 . 6( log Y 2 -log Y 1) JI= 1 l - - -~ ~ Van 't Hoff QlO X Centi5rade ~ -.)C2 y y1 10 010 = (-) xl-x2 y2 log QlO log y1-log y2 = 10( ) ~-~ D. Vi. Thompson x(also ~1) t Centit;rC~.de n v v t+n : _ _n v-x--t log V lot; X = t+n-log Vt n w \.0
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
40
Em.piriaal equations that have been used to express the relationship
of temper&ture to speed of development include the hyperbola, oatenary
and logistic. The widely used temperature aunmation rule ia the equation
r4 an equilateral hyperbola,
yx. k
or y(x-a) • k,
where 1. is tim., of development, ! is temperature, a is threshold tempera-
ture and k the temperature sunmation constant.
This rule states t/l..a.t the product of tilne by temperature above the
threshold is constant regardless of incubation temperature, and that the
1 reciprocal curve, - : Ia, is a straight line. Usually the observed re-y
oiprooal T&lues fall on a straight line only in the median portion of
the temperature range of development, and often the temperature-time
ourve has an exponential form, the reciprocal curve being o-ahaped. For
this reason Davidson (1944) bel16Ved that the temperature-summation theory
is an unsatisfactory representation of the faots and that its use should
be discontinued.
In 1926 Belehradek proposed the formula
a y • ::b
X
or Y: ~
X
a& a better method th&n ~ or Qlo for describing rate of development. The
temperature summation rule is of the same general form but with b • 1.
When it was neoeuary to introduce biological aero into the formula, the
equation be~e y • k 0 1 but since Bslehradek measured temperatures
(x-a)
in degrees centigrade and biologioal zero was praotioally 0°C for the
•k will be substituted for the a ueed by Belehradek in order to be oon-
sistsnt with •ymbols used above:
• j
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
41
animals studied, "it was not necessary to ocmplice.ta the formula by a
· !'itth !'actor" {Balehr~dek. 1929).
By logarithmic transformation Belahradek' s equation beoCJne3 log y
s log lc ""' b log x. which 1!1lJ111ea that when the logarithm of time is
plotted ag&inat the logarithm ot temperature the valuea lie in a straight
line. Belehradek found this to be truo for oonduotion in the soiatio
nerve of the fi-og, looCII\otion of .Pa.riUileoi um. a.nd embryological develop-
ment of the Mediterranean flour moth. Also, values of b were found to
be more constant than )l or QlOo
The data used by Belehr~ek for the Mediterranean flot~ moth were
frCII\ a paper by Janisch. who used the same data to dw.onstrate that tho
time-temperature relationship oa.n best be expres3ed by a catgnary ou.rve,
y • ; (ax+&. -x ) •
In this equation z represents the time required for developr.ent at the
given temperature! in def;rees oentigradaJ E! is the time for development
at the optimum tempera.tureJ .! iR a constant.
J&nisoh (1925) believed that the catenary fitted the observed data
throughout the temperature range of development and was later supported
in thia view by Uva.rov ( l9Sl). However" in a later experiment on the
rate or embryological development of the same moth. Ephestia kuhniella.
Voute 1n l9S6 obtained reaults that were oonaiderably different fran
tho•e obtained by Janiach. For points at temperatures above the peak
Voute believed that the catenary does not fit (O&vidaon. 1944).
A t~ ot the logistic ourve was found by Davidson to be a better
fit to the flour moth data than either the oa.tena.ry or Belehradek' a modi-
fication of the hyperbola. Davidson observed that often the temperature-
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
42
tilne curve was of the exponential form and the reoiprooal was similar to
a form of the logistic ourTe developed by Pearl and Reed (1920). This
formula,
has been suoaesafully applied by Davidson to describe the relationship
between temperature and rate of developm9nt at constant tenperatures for
six apeoiea of insects. In thia formula 1/y ia velocity. that is, r9oip-
rocal of the time requi.red to complete development at a given temperature
~~ L, a and b are constants. L ie the paraMeter representing the dis-
tanoe between the upper and lower asymptotes of the lo~istio ourve and
oan be calculated from the followin~ formula,
L: IP1P2PJ-P2 2 (P1+P3)
P1Ps·Pz 2
where P1• P2 and P~ are valuea for 1/y on the curve at three equally
apaoed temperatures on the absoiaaa.
Replacing 1/y with. P, the original equation 0111 be transformed to
the eqU&tion of & straight line. log8 ~ • a-bx. and the constants a
p
and ~ can be oa.loulated by the method of "least squarfts." In essence
this equation etatea that for a given set of data to be expressed by the
logistio ourve. a plot of the logarithm of L-P/P and temperature should
be points on a straight line.
(b) Fish. The early history of the searoh for a satisfactory law
relating taoperature to speed of development was centered around the
tsmperature summation rule, although it was uot identified as suoh.
•In the original equation Davidson uaod the symbol K. but to avoitl ooz•-
fuaion with k in the temperat\D'e &'I.IDmation rule, the K in Davidson's
equation wilT be replaoed with L.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
4S
Reaumer in 1735 suggested that the sums of the daily temperatures were
related to the maturation ot plantso Bonnet with ohloka and deCandolle
With plants were othera who early recognised the dependence of develop-
memt upon temperature. but it was a century later before quantitative
Observations of the effeot ot temperature upon deYelopment of fish eggs
were made (Thanpaon,. 1952). Hayes ( 1949) rev!Mel these early observe.-
tiona as follawwt
Davy in 1856 and Coate in 1858 gave eane fragmentary
figures showing that warmed water • .,.eels up -t.he deve1·
opinent of salmon eggs. Probably the first modern
work was done in 1859 by stephen H. Ainsworth who
experimented with egga of the brook trout,. Salvelinus
fontinalla,. in a little apr1ng fed fish pond near
Wea'€ Blocnfield. New York. Hil!! table showing the
inoubation periods of egga at varioua temperatures
was published by Norris in 1868 •••• Seth Green (1870)
stated thAt "trout eggs will ha.toh in 50 days at a
mean water temperature of 50°F and f"or ea.oh degree
oold.er or warmer five days more or lese will be re-
quired. the differenoo. haweTer,. increasing the rar-
ther we reoede f"ran 60 degrees."
W&l1ioh in 1901 suggested a ther.mal unit systsm and Apstein in 1909,
a temperature unit called "Tagesgrade." day degrees,. both being expres-
aions of the temperature summation rule. By Wallioh's definition a
temperature unit meant On$ degree above 32°F for a period of 24 hours.
For the chinook salmon frcm the Sacramento River reared at average tem-
peratures of' 43° to 60~ the number of thermal units to hatching we.s
constant at a.bout 900o
The "Tageegrade" is the product or temperature in centigrade units
and days. but differs :t'ran Wallioh' a idea. of thermal units in that the
threshold temperature was reokon•d frcm the lowest point at whioh devel-
opment oould take plaoe, rather than fran the free&ing point or water~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
44
Raiblsoh ( 1902) oa.lcula.tdd the threshold temperat,ze !'rcrn obser.,.;-s.-
t.i on a of Dannevic: ( 1895) upon t;he 1 nfluanc<:! o~ tootpera.tur3 on t:-ti'J dew1l-
opment of tho tl.:gs of the glaioe and the •nd. ';he data for incubation
temperatures a.:1d days to hatching were ca;1binad in pRirs to form aqua-
tions o.f the type (t1-x)n1 : (tz-x)n2 , in w:·.1ch _.!: is tempera.turt:J o~
l:louba.tL:m, x is threshold tem;>era.ture &.nd ::-. is nwnber of days to hatch---
ln.~. This equation was solved for x s.nd the average value for all the
pairoJd tem,?<1raturea wa.s Cl'nsid~red to be t!H3 threshold ternpara.t,u·e for
the species. Fo:r the pla.ice t':1A avera.ce was -2 .4°C and t._.e ranr;e -L2°
to -4 .oocJ for the cod the av9rage wn.s -3 .~°C and the runge ·1.2° to
-13.2°C. !.!sing t;he aver&.r:;e values, Reibisc:.,• a calcn.J.1~:itions were constant
and for this reason ne cvncluded thAt the t.r~ory of tcmp~ratu.re summation
with the proper tem?era.tl.rro threshold was va.Hd.
Johansen and Krogh·· ( l'Jl1) took exce~,tiun to tha ill ea. of' P..eibisch
thAt a certain amount of heat or energy from outside of the egg was nee-
essary for development.
When the eggs ha.v~ the same temperature as their sur-
roundin6s, they get no SU??lY of heat from out5ide •••.
The energy whioh ia undoubtedly neoessary for the
develop:nent, is darlvoo in the oase or fish e,::;~s~ a&
in all other e~gs, from the ohamioal processes involved
in the metabolism of the agr;.s., •• The tsup !3ratw-e must
be looked upon aa a faotor which will have a certain
influence upon the velocity of the ohem.ieal reactious
and other processes involved in the development. The
theoretical problem is to obtain a quantitative meas-
urement of this influence and to express it in such
t.:~rms that a canpari son with regulR..r ohemi oa.l reae-
tiona becomes possible.
Using De.nnevig' s data. Johansen and Krogh found QlO also to be unsat·
isfactory but believed the temperature-development relationship was linea1
when temperature and the reoiprocal of time to hatching were the variable 1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
45
This means that the change in rate of development is proportional to the
ohanga in temperature, and the equation that expresses this relationship
is the temperatura s~~~tion rule in its reciprocal form, t ~ ~ , in
which y is time, x is temperature and k is the temperature summation
constant.•
A variation of the ~eneral form of the Bslshradek formula, y : ~ ,
X
was used by Price (1940) to describe the rate of development of the
whitefish, Coragonus ~upeaformis (Mitohill). For the Lake Erie white-
fish S?awnin~ bebins in late Novamb~r when decreasing water temperatures
a?proach 6°C and the four-L-onth incubation period is at temperatures onl:
sli~~htly a.bove freazinr. rrir.e found that the rate of development was
different above and below 6°C and proposed a two-part equation to descril
this condition. Hie equation is of the form
T -M l..
• Alt J
where T is time of developman~ and t is tamperature. Values of A1 aver-
aced 1.13 and ~f A2, 1.13.
For the Salmor~idae the hi story of exper ~ants in whioh there are
some data relative to the rat"' of development is summarized in Tabla a.
These experiments were reviewed for formulae ex~ressing the temperature-
development rel~tionship and if none was given, the ourva of temperature,
!• and reciprocal of ttme. 1/y, was plotted rram the data. This curve
was arbitrarily selected as it was simple to plot and as likely as any
to have a linear relationship. The slopes of these curves increase with
•Nomenclature varies with authors but for consistency in this report
translation to these standard terms wiJl be made where necessary.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.TABLE 8 Su..amary of Temperature-Rate ot Developoent .&x.perimanta with Salmoni.dae lggs Range of Temp. Source Fit of tllt curv• I71-=X/kt.o-the relationahip Jnveatigator Date Species Temp. l'attem o_f Data ____ _Q{ _ _t•PAratu.re t.9J~t~LC>! cle'YelopJDBnt Ainsworth 1868 brook 37-54°F x* bl Poor; !or 50°F, 1/y is high Green 1870 trout No data; incubation perioc:l is 50 ~s at 5QOF Wallich 1901 chinook 42-510y X h Good; suggested te.uperatu.re unit s;p t.e.m Kawajirl l9Z7 masou 6-16°C c*** ell# Good; h.igh mortal.itiu; mn-circulating water Kawajiri 19;(S rainbow 7-l2°C c,x e Fair Gr~ 1928 brown 3-l~C X h Poor; for temp. above 5°C, 1/y ia high Belding, et al. 1932 A. Salmen 3.3-42 F X h Very poor; points widely scattered Emboey 1934 brown 2.-ll°C c,x e,h foor; similar to the results of Gray (1928) Embod_r 19.34 brook 2-140C c,x e,h Fair; above ~C, 1/ y values aro high Embo~ n rainbow 3-l6°C c,x e,b Good; Embody II lake 2.-lQDc c,x e,h Fair; ccnca.ve to absciesa Merriman 1935 cutthroat 6-ll°C c e Good; only 3 pointe Rucker 1937 sockeye 8-l4oC c e Go<Xi; only 3 points Foster 1949 rainbow 43-5:f>F X El Fair; (.x-ray experiment) Donal dam 1950 sockeye 55-J:i'F X e No data.; eggs moved to )20F at various stages Donalda on 1955 chinook 55-b'f>F X • Fair; egga moved to 5h fran high temperatures Burrows 1956 chinook 35-60°F c • Good; includes la. tem,Ferature data *creek trout, Salve1inua fontin&l.is; chinook, Oncort:gnchua tshamscha; masou, Oncorilynchua .!Da80Ui rainbow, Salmo gairdnerii; brown trout, Sal.mo trutta; Atlant.:ic salmon, S&lmo salar; lake trout, Cristivomer na.maycush; cut-throat, .::ialmo claridi; sockeye, Oncor}Vnchua nerka. **x-changing temperatures ***c-constant temperatures llh-data from hatchery records #He-experimental data &:
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
47
temperature and in general are slightly s-ahaped when the experimental
temperatures extend O'V!'r the entire range of temperatures at whioh devel-
opment 1e possible. In several experiments the relation of 1/y to ~ was
not lin•r• Rockwell (1956) has plotted the rate of development curves
tor forty-one experiments with tish including the experiments listed in
Table 8, except for the experiments ot Foster ( 1949) and Burrows ( 1956).
In addition to the rate of development experiments listed in Table B.
other temperature experiments with chinook salmon have been carried on
and include the follmrlnga Brett ( 1952) d~ennined the upper and lower
temperature tolerance for fingerlings of fiTe species of Paoifio salmonJ
Johnson and Brice ( 1953) made observations on the effect of water tsnper-
ature during incubation on the mortalitl of chinook salmonJ Donaldson
(1955) reported on the survival of the early stages of the chinook salmon
a.rter varying exposures to upper lethal temperatureaJ Olson and Foster
(1956) determined the temperature tolera~oe of er,es and young chinook
salmDn at temperatures abave and below that of the Columbia RiverJ Bur-
rows' 1956 data are not published, but incblde in part observations on
mortality and ~ .2.£ develoeent of chinook salmon eg~s and fry at low
tsnp'51"aturea.
Controlled temperature experiments with Pacific salmon other than
the chinook include those of Kawajiri (l927a), Ruoker (1937), Donaldson
and Foster (194~ Donaldson (1950) and Rockwell (1956).
Constant temperature experiments. Other than Wa.11ioh' s temperature
summation ~stem and a provisional velocity of development curve for
Pacific salmon eggs by Rockwall (1956). the tamperature and rate of devel-
opment relationship for chinook salmon had not been determined. In the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
48
eearcb to~ an equation that would beat describe this relationship esti-
mates of l'• QJ.o and x were made and the fit of t:he data. t1' the temperature
summation ~ule. the hyperbola and the logistic was tried.
The incubation tempe~a.tures and the number of days to hatching for
Experiments I and II are pres•nted in Table 9 and Figure 1 o The data
uaed in the searoh for an equation to describe the temperature and rate
or development relationship were selected tram lots reared at tempera-
tures between 39.8° and 57 .sor. Both above and below thie range mortality
inorea .. d markedly (Fig. 20) and the rate of development ourve flattened
(Fig. 15). The rapid increase in m~lity is interpreted to mean that
the increase in deaths is due to temperature, the flattening of' the rate
of development ourTe oould b8 interpreted to mean that the fast-growing
individuals are killed first at high temperatures and the slow-growing
individuals are killed first at law temperatures. To avoid the possibil-
ity of the influence of let.~l tamporaturos upon rate of development th~
data. were limited to thosu lots reared in the temperature range 6 39o8°
to 57 o8'?. There were only four lots from eaoh of Experiments I and li
that were reared at oon&tant temperatures in th.is range and therefore
the data .fran th8 two eJqJeriments wera oanbined.
In oanbining the results of EXperiments 1 and II it is assumed that
bstween broods of dif.fereut years the rate or development is not statis-
tioally significant. Differences indicated by the rate of development
trend lines for Experiments r. II and III in Figure ~ are not groat.
The relatively gJ"•ater deviation of the trend lines at high temper-
atures (Fig. 8) may be expected for two reasons. First. error in esti-
mation of hatching time was slightly greater when the incubation period
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
49
TABLE 9
Wc1ter 'Temperature, .lncubot icn Feriod and F~r Cent Mortality of
Green River Chiuook .:>o.l.non Eggs in Ex.!Jtr.ir~nts I o.nd ll
Temperature
I·c.ttern
Constant
Constant
G~an~.i.r~;
at city
w.:::..t--:;r
te:n;~ rat ure
Changing;
at city
water
tempt! rat u re
Year Lot
1951
195.2
1951
1952
l
2*
3~
)*
l*
7
10
9
l*
7
'i
1~..o
BA
8C
4
8
8B
-
Tem~era ture
Cf OC
4~ .. 7
5C..t
55.1
60.2
t2.4
l4.b
45.~
5\... • ..2
:;4.6
CJ~.o
._.7.4
46.9
47.0
4'/ . .G
58.5
59.0
II
II
l.ll
4.Jj
7.v6
10.33
l~ .. SJ
15.67
l6. 89
1:3.11
7.JJ
10.11
12. 'j[
l4.j)
1) .44
lb.07
ld.2.2
19.4 ...
i~. -16
8. -~8
3.33
8.44
14.72
15.00
11
O~ys to .. Fer-Cent
5v% Hatch Mortality
128.6
'19.1
)l • .2
4J.O
31.. J
,...,1.4
.:::8 • .)
5G. '1
y'.8
54.0
j~.1
-• I
o:.::..j
06.1
r. 5 .o
b2.5
32.4
J2.. 7
II
11
1UO
0
13 ,.
.)
22
78
99
1
.!.
2
2
35
C3)
10v
luC
15
5
2
j
4
* Selected for curve fittin5 because of l0\'1 ,nortalitj·
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
0
100-
~
0
~
~ at
() so'~ "-0
~
0 0
'1..!\
0 ..p
Ill 6o~ ~
A
..... 0
M co
(D
l 4oi~ oo ,, 0
0 0
0 cP
0
2oll
II
II J_,
. l .· .:-.1,-=
40 45 50 55 60 65
Average Temperature, OF
Fieure 7. Averaee Temperature and Number of Days to 5o Per Cent Hatch for
All Lots Reared at Constant Temperatures
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
).0 -
::r: u
~
~ 2oS -
(.)
~
0
U'\
0
E-4 2.0 -
C/}
~ ~
~
&l
I 1.S
~
I
I
I .·
I .. ··
I /
I /
I .··
I /
1 .. ··
__ 1_ .•.• ··-·--·-··· .. L-------···
/
/
55
AVERAGE TEMPmATURE, <7
/
/
/
p
/
/
.):>
II,,.
y
L
60
--~
6)
Figure 8o The Average Temperature and Rate of Development for Eggs of the
Chinook Salmon from Green River for Experiments I, II and III
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
62
was short (page 27). Secondly, the ordinate of the rate of development
curve is the reciprocal of the number of days to hatohing, and the graph-
ioal representation of one day for a short incubation period, that is.
high temperature. is greater than for one day of a long incubation per-
iod. For example • the di.t' terence in 100/y for y : 29 and y 11 ~o is 0.12 J
and tor y ~ 100 and y • 101 is OoOl.
(a) T~perature coe.t'tioisnta. Using the eight lots from Table 9
that were selected for low mortality rates a.nd combining them two at a
t1me. 28 o~binations were obtained for which ~. Qlo and x were calcula-
ted (Table 10). Omitting the values when the tsmperature difference is
leas than 0.3°C•. the range for~ was 12.000 to 29,500i for QlO• 2.11 to
6.40J and for x, loOB to 1.20, respectively, clearly indicating that
these coefficients are not constant for the relationship of temperature
to the rate of development for chinook salmon. The average value for ~
of 20,000 and for Qlo of 3.64 agrees with the atatement by Hayes ( 1949)
that "any Qlo value for salmon and trout can be converted to the oorres-
ponding value of Arrheniua' fo~ula with negligible error (5 per cent)
if multiplied by 5600~"
Since the values of ~ and Qlo are not constant, then the relation-
ship between the logarithm of the speed of development and temperature
is not linear (page SB). To investigate the shape of tho curve expres-
sing this relationahip two graphs were madeJ for ~ the variables were
the reoiprooal of temperature in Kelvin units and the logarithm of the
•For alight changes in temperature the relationship of temperature to
tbne of inoubation is not accurate due to experimental error and occa-
sionally may shaw a slight increase in incubation time with an increase
in tcperature o
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
TABLE 10
Temperature Coefficients and Threahold Tcperatlll"'es for Chinook Salmon
from Green River Reared. at Conatant Temperatures
Temp., oc• Temperature Coe !.ficienta Threshold Tem.!Jerature, !*
:.tlo* x* 1i OF oc Xl ~ u
4-33 ?.06 5.93 1.19 27700 32.0 0.00
" 7.33 6.40 1.20 29500 .32.6 0.33
" lO.ll 4.98 1.17 25400 33.0 0 • .56
" 10.33 4.83 1.17 21.,?00 32.9 0.50
If 12.56 4-JO 1.16 2.3200 33-4 0.78
n 12Q83 3.96 1.15 21800 )2.9 o. 50
II 14.33 3.79 1.14 21.300 3.3.3 0.?2
7.06 7.3.3 (1.3.2) ( 1.29) (49800) (.38.2) (.3.44)
" 10.11 4.26 1.16 2.3200 34.8 1..56
II 10.33 4.05 1.15 22200 34.5 1.39
If 1.2.56 ,3.67 1.14 2<J1()0 35.2 1.78
11 12.83 .3.27 1.13 18900 34.1 1.17
" 14.33 3.20 1.12 18700 34.8 1.56
7.33 10.11 .3.79 1.14 2)900 33.9 1.06
" 10.3.3 3.61 1.14 20000 .3J.5 0.83
" 12.56 3.42 1.13 19600 34.7 1.50
II 12.83 3.0.3 1.12 17600 3.3.4 0.78
If l4o33 3.02 1.12 17?00 34.3 1.28
10.11 10.33 (1.93) (1.07) (9200) (21.4) (-5.89)
If 12.56 3o~ 1.12 18000 .36.1 2.28
" 12.83 2.42 1.09 14200 32.2 O.ll
u 14.33 2.61 1.10 15500 34.9 1.61
10.33 12.56 3.18 1.12 19100 J7 .o 2. 78 .. 12.83 2.47 1.09 14600 32.9 0.50
II 14.33 2.65 1.10 15900 .35. 5 1.94
12.56 12.83 Y 2 -Y1 1.t negativ •
" 14 • .33 2.11 1.08 l;.;ooo 31.9 -0.06
12.83 14 • .33 2.98 1.12 18000 39.8 4.33
Average 3.64 1.13 20000 34.1 1.17
*See following page
( ) ~ -x1 less than 0.3°C; estimates inaccurate in this range
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Oy
39.8
44.7
45.2
50.2
5().6
54.6
55.1
57.8
TABlE 10, continued
y
oc da..v!
4.33 128.6 x = temperature of incubation; !or u
temperature in degrees absolute
7.06 79.1 = 273.18 oc
7.33 73.4 y = nUIIber or days to 50% hatch
10.11 50.9
a = threshold temperature
10.3.3 50.2
12.56 38.8
12.83 40.0
14.3.3 34.0
1 1 (4.6) 1og(i)2 -1og(y)l
u = 1 l
~-i2
log Q10 =
(10) log(;)2 -log (;)1
~-X1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
55
1peed of developmentJ for 'ho the Tariables were temperature in centi-
grade units and the lo~arithm of the speed of development. From inspeo-
tion of Figure 9 it would appear that s.6°C h a oritioal temperature.
Values of \1 and Qlo above and below this temperature are as followsa
Coetfioiant "' QlO
4.ow -8.50C 28600 6 .~2
8 .s \) -14 .J°C 16100 2.69
4:.Jo -I.;..o 0 c 211!00 S-9~
The values for~ (or Q1 ) in Table 10 may appee.r to be relatively
oonzt&nt, but x is one-tenth the log of Qlo and therefore no better a
measure of the rate of development tr•n QlO• the reason for the more
constant 'ftlue of x is that the number is obtained from that part of the
log table in whioh a large ohange in the logarit~~ corresponds with a
relatively snall ohange in the number. Thompson (1962) lists the~ or
Q1 for a great variety of organisms and points out the oonstanoy of val-
ue~, from 1.08 to 1.20. However, in terms of Qlo the range of values
for the same data is 2.2 to 6.2.
In oonolusion, a single value for any one of the thermal ooeffi-
oienta, ~· Qlo or x, does not adequately describe the rate of development
of the ohinook ~lmon egg.
(b) Threshold temperature. The first step in fitting the chinook
salmon data to either the temperatura eummation rule or the Belehradek
equation was to estimate the threshold temperature, also oalled "schwelle,n
biological~ or ori tical ~~growth. The threshold temperature
is ~ faotor in both of these equations and often has been disregarded
when the incubation temperatures h~ve been measured in centigrade units.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
56
0......._
..._
-0 l.ll b-..._
r.:... A 0 ........... ...._ ..._
~5
li
--oQ "p. I! ~~ ..... ' -_,
'~ II -1.2j~ ..,._ go , .....
' ' .-i ' -~ ·o
'
f:;o CA..
' ' m'V\
1.0 .~ ' ' ~~ ' ' ' ~ ' §~ ' ' ' '0
OoB --~--~~==-~~---_:____~-~-~-=-=-==----
.003475 500 525 550 575 600
AVERAGE TFl'IT'ERATURE, 0 ABSOUJTE
!! .0 " / II /
I il /
l.q r-06
/ I:=-. il /
0 / ,.......
:I
/
~6 /
/
~~ / /
p:Y
// B
F-• ,I _..// '-.E-t I
§~ :r // 1.2 !i // "Q"
/ 10
r-i I /
-~ /
~ ~
0 1.0~~ /
~'V\ /
/ f:;o /
~~ /
/
/ §~ /
/
II
/
d'
o.all ~-L
4 6 8 10 12 l1 15
AVERAGE l»U'ERATURE, 0 c
Figure 9. ReJAticmship of Te:rrq>erature to the Logarithm of 1000/Number
of Days to Hatchin;·. A, Temperature aa the Reciprocal of the
Absolute Temperatureo B, Temperature in Degrees Centigrade
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
67
The threshold temperature of many animals is near 0°C and therefore the
oorreotion for this faotor 1 s not great.
Reibiaoh estimated the threshold temperature by a method already
described (page 4S)J Johanaon and Krogh (1914) extrapolated the rate-
temperature curve to the ! (temperature) axia and called the point or
intersection the threshold temperature, but acknowledged that "it is not
legittmate to assume that the curve of development remains straight beyond
that part whioh has actually been inv~stigated." Krogh {1914), Shelford
(1927) and others have recognized the change in rate of development at
high and low temperatures, but an estimate of threshold tamperature other
than by extrapolation or by the method of Reibisoh has not been proposed.
TWo other eatimateG of the threshold temperature can be derived frcm
the temperature sUJml&tion equation, y(x-a) : k:, where y is the number of
days to hatching at temperature =.• ~ is the threshold temperature and k
is the temperature summation constant. In one method the number of tam-
perature units to hatohing ie assumed to be the same for eggs incubated
at one temperature as at any other temperature. Thia oan be stated in
equation form as follow• a
Yl(xl-e.) = lc
yz(x2-a) • k:
Yl(xl-a) : Yz(x2-a)
Reibisoh calculated the threshold temperature in essentially the same
manner for paired observations and averaged the values to determine the
threshold value for the species. For the chinook aalmon the value
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
58
oaloulated in this manner was M.l°F (Table 10).
The general estimate of the threshold temperature for the epeoiea_
u1ing the aame data. ia the regre1sion ooeffioient ot x1Yl-X2Y2 on Yl-Y2
whl.oh waa 32.8°1 tor the ohinook l&lmon reared at oonstant temperatures.
The ~lue of the regresaion ooeffioient is a better estimate of thresh-
old temper&ture than the average of the paired observations for two
reasonsa (l) the better fit to the data as ehawn in Figure 10, and (2)
the. smaller value for the ooeffioient of variation of k when 32.8°F rather
than 34.10f was used as the threshold tamperature. The ooeffioient of
vari~tion~ c, in per oent w~s 3.47 and 4.25, respeotiv9ly~
A seoond method of eetimatinr, threshold temperature involves both
a and k. wher·;;~!s k was eliminated from the above equation. Using; the
~e symbols, the temperature oonatant and threshold temperature are de-
rived as follawse
1 • x-e.
y T
~ = -~)+(i)(x)
Thia equation is in standard form for the equation of a straiEht line,
y : a+bx, and both (~) and -(~) oan be determined. The faotor ~
ie the regrea1ion coefficient of the rate of dgvelopruent, ~. on tempera-y
tur~ x. By substitution in the IJt&nde.rd form the threshold temperature.
a, is found as follaw5a -
.. ~ = (;)-(~)(X)
a. :X T) k • 'f -<-y
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
' 3~
loot
0 ---
0
1
Jo<t
i
l
1.ar-
/
/
/
/
/
/ ••
/
/
/ •
/
/
a -:: 34o1 °F, from Table 10
/
/ ••
20 ho
40
\ /
/
/
// ' / a = rer,ressian coefficient
/ /~ o.f (~y1-~y2) on (y1-y2) = 32.JoF
45
. .L ---__________ .._ --··· • --
Bo 100
a = x-k(l.) = J4.0°F y
1 = ree. coefo of the
'k ar development on
temperature
50 55
rate
AVERAGE T:El-IPERATURE, 'T
Fieure 10. Three Estimates of the Threshold Temperature, a, for Eight Lots
af Rlcperiment I and II. A, .! Determined from Values of ~ and z.
B, .! Determined from Values of ~' z and ~·
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
60
-("I a • x•k -) y
By th:..s method the threshold temperature for the chinook salmon was foWld
to be ~3~8°F. The linearity of the relationship of the rate of develop-
ment to temperature ia shown by the closeness of the points to the
regression line in Figure 10.
There is a good fit of the data to both of these two new methods of
estimating the threshold temperat~·e. The latter method is preferred
because the regression lir~ is not required to pass through the origin
-T a.r.d the variables x and -a;;d the oonstant k are more easily detennined.
'./ -
The ~onfidenoe li;ni ts for the thrdshold temperature derived by the
second method can be det~rm1ned ly solving the following-equation for as
t t. A \ .L (a.. -!:}___ .,\
n -.2. .!_ 1-["' + - - - . 'j 1_ l ,_L -}1-) j
t o I
estimate 1 1 ot 1C : "iC as defined above
estimate of a ~ • ~ as k defined above
tn-2 • 5 per oent point of Student•s t for n-2 items
s1•x • standard deviation from re&reasion of the rate
y of development on temperature
.x • mean of temperaturo observations
n : number of observations
The confidence interval for the threshol~ te'!T;perature is defined b~r the
values of the two roots of the equation., and for the chir.ook salmon data
these values were oaloulated to be ~3 .. 0° to 3-fi.6°F.
•Equation derived by D. G. Chapman, Mathemntios Department, University of
Yiashia~?.:tor ...
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
61
In conclusion, the threshold temperature for Green River chinook
a.lmon reared at oonatant tcperatures in the range of 39.8° to 67 .8°F
ia about s:s.sora values ot thJ"•• different estimates were !2.8°, 33.8°
and 84.loi'. The 95 per oent confidence internl waa 33.0° to 34.60f for
the th.J"eshold temperature of ~!.80, o
12) Temperature ll\lml!ation rule. The threshold tmperature having
been estimated, the proceaa of curve fitting waa restmled,. The linearity
of the regression of x1y1 -szy2 on Yl-Y2 and of 1/y on~ in Figure 10
shows that the temperature IIUI!ID&tion rule is a good expression of the
relationship between the rate of development and temperature for inouba-
tion of chinook aa~on eggs fram Green River in the temperature range of
39o8° to 67 .eO,.
{d) ~lahr~ek equation. The N.t of the ohinook salmon data to the
Belehradek equation and to the logistic was also tried. The Belehradek
equation by logarithmic transformation beoc:mes the standard form of the
equation of a straight linea
k
y • ::b
X
log y : log k-b(log x)
Tlut value of b, which is oalled a thermio coefficient, is the regression -
coefficient of log y on log x. For the ohinook salmon b was -oo968 when
corrected for a threshold temperature of ~3.8°F and -l.l2_for the unoor-
rected data (Fig. 11). The value of~ for the embryonic development of
Salmo fario aa calculated by B~lehradek (1929) was 0.991 for Onoorhynohus
nerka as calculated by Rucker (1937), 1.06.
In aonoluaion, since the t~parature summation rule and the Beleh-
I
n.dek equation are identical when b • 1 (the chinook ciata ahow the value
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
2.1-
0.2 -
1.8 -
62
o',
.......
........
........
.......
........
........
........
.......
.......
.......
.......
........
'O 'a
........
........
........
........
........
.......
' ' Belel'lnidek '
.60 .70 .Bo .90
LOJ ~TORE, c
'ao
........
.......
........
........
.......
• .H.
'o:> .......
.......
1.1
~ B
.......
........
........
........
........
........
........
'0 'O
' ........
' ........
........
........
........
'ao .....
........
........
U:>gistic ' ,
4 6 8 10
TEMPERATURE, °C
.........
'c9
.........
]2
........
.........
........
'"'0
1-
ili
........
'o
F'igul"e llo Rel.atianship of the Belehrcidek and Logistic Equations to the
Rate of Deve1opnent Data for Eight Lots of Experiments I and II.
A. Relationship o.i' the Logarithm of Temperature to V1e
Logari tbm of the Number of Days to 50 Per Cent Hatch.
B, Relationship of Temperat.ure to the Logarithm of L-P/P.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
63
ot ~to be ~pproximately 1), then the relationship of either (l) the
speed of development to the temperature or (2) the logarithm of the num-
ber ot days to hatehi~, to tho logarithm of' the temperature is linear
for lote reared at oonetant temperature in the range from Z9o8° to 57.8°F·
(e) Logi.tio curve. The logistic ounre uaed by D&vidson ( 1944) was
ot the farm
Replacing l/y with p the equation ia developed as follawsa
p : L
l+ei-bi
L-P loge P:: a-bx
The last torm 1a again a form of the equation of a straight line for
which ~ and ~ can bo determined by standard methods. L oan be estimated
aa described on page 42. The relatiozahip ot log .k:!. to x should be lin-p -
ear it the logistic e~uation describes tho t~perature development rela-
tionahipJ for the chinook salmon it appears to be ao (Fig. 11). 'fhe
'ftlues for the oonstanta were :5.96 for L, 2.46 for .!• and 0.242 f'or b,
the equation for the curve being
In Figure 12 the lo~istic ourva closely ~its tho poir.t& that are
determined by the relationship of temperature "t.o the number of days to
hatchingJ and the reoiprooal of the logistio ourve fits squally well to
the points that are determined by the relationship of temperature to the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
>..
II
:I:
(.)
8
E2
E-t ~
0
~
0
lJ\
0
E-<
~ A
f:>
&1
i
... •
1751-)350
/
/
I \ /
\ /
/
\ /
150· \ / -300
\ 0 /
\
\
\
125-~1 -250:.:. :n
100.
7.-' . ;)
so-
25-
/
/
/
/
40 45
1+
y.:.
---_!
5o
TEMPERATURE, <T
y
.03951
2.459-.,242x .. 200 c
~150
I
I
~ 100
-----
-50
1 2oh59-.2L2:x -...e
3.957
_, ~------~_j_ -~--~-d 0
55 60 63
Figure 12. The Relationship of Temperature and the Number of Days
to Hatchinr: , to the Logistic Curve and Its Reciprocal
for Eight Lots of Exper:i.ments I and II
II
~ Q
~
~ s
§
~
0
~ p..
0-.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
65
speed a£ developnent (the reciprooal of the number of days to hatching).
The relationship of tamperatw-e to speed of development was shown to be
linear in Figure 10 and to fit the logistio in Figure 12, but this is
explained by the faot that the temperature range of the lots aeleoted
for ourve fitting lies in the region of the point of infleotion of the
logistic curve, a region where the logistic ourve is nearly a straight
line.
The apparent good fit of the logistio ourve to the points in Figure
12 does not necessarily mean that the logistic expressea the theoretical
relationship between temperature and rate of development. For three
ourves in whioh the deviati)nB of the points from the curves were less
than ahawn here for the ohinook salmon, Brawninr (1952) tested the good-
' ness of fit by the X test and found that the probability of the calou-
lated curve describing ~he relationship of temperature to the rate of
develot:ment was less than 0.0001.
In oonolusion, for the temperature range of 39.8° to 57.80p the data
fit the logistic curve, but the fit is no better than either.the t~per-
ature summation rule or the Belehradek eq1~tion.
Altered temperature experiments. Experiment III differed fran I
and II in that eggs from four raoes• were used and the temperature pat-
tern --.a ohanging rather than constant. The temperature history for eaoh
lot identified by the SLme symbol waa similar. that is. the temperature
history of the eggs of Skagit Lot 1. Enti&t Lot 1, Green Lot 1 and Sao-
ramento Lot 1 was similar. Water temperature. inoub&tion period and per
ot!The ohinook u.lmon from the Slcag;it, Entiat, Green ana Sacramento Rivers
are considered to be separate raoes.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
66
oant hatoh are given in Table 11.
(a) Incubation rate. The rate of development of the Sacramento eggs
was fastest, with Green, Skagit and Entiat eggs following 1n the order
ginr1e To determine the rate of develo~ent for ee.oh race relative to
the raoe from the Sacramento Ri 'ftr, the n~r of ~s to hatching for
eaoh raoe was divided by the corresponding value for the Sacramento River
raoe. Thia was done for eaoh temperature lot by races and then the aver-
a.ge 'ftlues for the four races were determined. For the agga from the
aalmon of tha Sao~anto, Green, Skagit and Entiat Rivers these values
were 100, 97.2, 94.3 and 92.,, respectively. Actually, the average tam-
peraturea for similar temperature lots varied slightly between races e.nd,
for comparative purposes. the number of days to 60 per cent hatch was
adjusted to a common temperature by linear interpolation (Table 12).
Evidence of ditferanoaa in incubation rates ot the four ra.oea was
obtained from three souroess ( 1) inapaoti on ot Figure 13.., which shows
the number ot d~s to hatching for each race at eaoh temperature, (2)
tast of the significance of the difference in the number of days to hatoh-
ing ~ student's !• and (3) the oonaistent ranking of the races in regard
to the nwnber of days to hatching at various temperatures as shown in
Figure 14.
The ralat1onahip ot the m.aber ot days a.t'ter the start of the exper-
iment to the cumulative percentage h&tohad is presented in Figure 13 and
ehaw• that the Sacramento agg lots completed hatching before the Entiat
egg lots began except 1n Lot 2. Lot 2 eggs were hatched at temperatures
of S4° to 36~ and coneequently the hatching period wa.a extended. The
5-96 percentile deviation, Pg5•P5, of the day• to 50 per cent hatoh
2
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
67
'I'h.EIE 11
Wo.ter Temperature, lnc ub~ tion t'er iod a:.d h;r Cc:n t 1ie;rtd ity
of the Chinook So.l" .. on Ec.:;gs of Ex f-er :L:c; n t Ill
Te;ni-. , F Nc. of I er Cent
Stock Lot 3tG1r'L Low End f..vc. Duys .l.!vrtality
.Skagit 1 45 34 45 j8.8 1.)~. (; L.
River 2 50 34 j6 39.8 112. \) J
J 55 44 44 4').4 50.C 2
4 60 51 51 55.9 38.7 2
5 6-) j') 59 61. J )1.) 40
b* 55 54 55 54.7 46.7 8
Entiat 1 45 34 46 33.8 1.53 .0 6
River 2 5U 34 ·y 4().6 lG-::.0 2
3 55 4L 42 48.4 62,2 2
4 6G 51 51 55.2 4~.3 2
5 65 58 58 bl. 3 33.5 11
6* 55 54 55 55 . .2 4C.4 9
Green 1 45 34 41. 39. () 125.0 42
hiver 2 50 34 36 41.6 97.6 J6
3 55 45 45 50.2 51.8 63
4 60 52 52 56.3 30 .l bj
5 65 63 63 64.0 100
6 56 54 54 55.4 39.7 52
Sacrc.u11ento l 45 34 4j 39.0 124.0 2
River 2 50 34 36 41.6 97.4 2
3 55 44 44 50.5 413.6 1
4 60 53 53 56.5 35. 5 1
5 65 60 GG b2. 2 LcL 4 21.
6 56 54 54 55.5 j8.0 3
* Incubated in water of low oxygen content
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Observed and .Adjusted Te:aperatures and Days to 50 Pt:!r Cent Hatch for
Ghinook Sal:n0n Eggs Reared at Cnanging Temperatures, Experiment III
Lot Stock ----
1 Sacramento
Green
Skagit
Entiat
2 S.;,.cramento
Green
Skagit
Entiat
3 St:.c rwiJ~::; nt o
Crreen
.:::ikaeit
i!;ntiat
4 Sacramento
Green
.::)kagit
Entiat
5 Sacramento
Green
Skagit
Entiat
:S* .::>acramento
Green
Skagit**
Entiat**
Observed
Ave.
Temp.
,oF
39.0
J9.0
38.8
)8.8
41.6
4}.6
J9.8
40.6
50.5
50 • .::
49.4
48.4
56.5
56.J
55.9
55.2
62.2
ol.J
61.3
55.5
55.4
54.7
55.2
Days
to 50%
Hatch
124.0
125.0
1J.2.0
13J.O
97.4
97.6
112.0
102.0
48.6
51.8
)t; .o
b2.2
35.5
38~1
j8.?
42.3
28.4
31.9
jj.5
38.0
J9.7
4r;. 7
46.4
Adjusted
Ave. Days
Temp. to 50%
,°F Hatch
39.0
II
II
II
4}.6
11
II
11
5U. 5
11
11
11
55.2
"
11
n
cl. 3
II
II
1;24.0
1.25. 0
128.G
130. (j
97.4
97.6
1Cl.4
96.~
48.6
51.2
5_;,.2
56.0
)8.2
40 . .)
40.5
42.j
31.9
jj.5
------------------
0 *"Controls", temperature range 56 -53 F
**Incubated in water of low oxygen content
5-95 fercent-Hatching
ile Deviation Rate Relative
to 5C% Ho.tch to Sacramento
).,27
4.31
1.85
;;,12
D.OO
lC.u8
5. 1.2
~-.12
2.75
--.98
2.CO
..::.12
l;. :)0
1.16
2. '/2
l.:C:j
1.80
2.bt
L,8J
100.
97.2
96.9
95.4
100.
99.8
96.1
101.
luO.
'14.9
71.4
86.8
lvO.
94.8
94.3
)0.j
100.
'12.8
88.4
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
3o5r
'i I
:I
:I
3.0 11
;:I:
u
E-i
5§
~ 2 5 I P'l • 1:
0
~
0
l.n
0
E-i 2.0 ;._
Cf)
~
z:§
J'x..
0
i
8
1.5 .:
...;
jl
II I.o ir
I ---Sacramento
-----Green
--·-----Skagit
--··-Entiat
0.5 -~---~~=--~~~----0 L~ .-_. __ ,~~~~-~ __ __l _ ·----=~·
4o h5 5o 55 60
AVERAGE TEMPERATURE, ~
Figure 13. The Average Temperature and Rate of Development for the
Eggs fran Four Races of Chinook Salmon in Experiment III
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.wr 2 U.6°F II:trl )9.00f 100'" .... ----·/"? ........ •• / ..... .. ' . ·I ,• . I I ~ ,./ / i / f../ i . t.) ,.. I :-1 : I : I wrs 61.30oj wr 4 55 .20p' wr 3 ;o.sor .. . . . : ~ : . : . : . . . . I ~ 7r! _ i I ! ,. i 1· 1 1. l rt:ll ~ = I. : = ~-. I. ! 1 ,. l 1 · ; ! ' ' I ' ( ' ' ' . I . . . . I . . . . . . . . . . . . ~ ! · 1! ! I r' : i· r.. . • I ' · · · · I o : . l I : . I : = ~ ;o ,. : 1 : : 1 : : · . . . . . . I . . . . . . < :1 r I : · J : ' ..... ~ : : . = I , : : . o u~ fJ' f f i , ( ! ij . . . { . . . . . I . . I ' ' ' I ' ' ' ~ 2S -f ~ f ! l . ,. / f I • . f . I . . . . . . ~ : ' : : .· . . i . . . . . J . . . . . : : . . . . . . ! I ! I ! I ;· / f· . . . . . . . . I u . I ) . _,1 . . .' I ' ' I ' . . . . . .. .... -,_. . . -.--, --·"f.·--· . ' . -.-. -1 .-:, ... , -' -.·--· 0a~ Jcl -~ 35--LO ---------5o !iS · • 90 --··· 100 • · --uo· --fw--DAYS APTER START OF EXPmlMEm --sacramento --·· -·Skagit Green -·-Entiat F"\17ll'r'A 1),_ r.nTrtn1o+:4v• p.,...,.A...,+ .. .-.• ,.I" .., ... _ l.T •• -\..---~~' .,. ___ TT--'--"---" •
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
71
(Table 12) is three to four tUnaa greater for Lot 2 than for any of the
other lots, and for this reaaon the 50 per oent hatching date is a less
reliable estima.t• or hatohlng to"r Lot 2 than for lot& with a shorter
ha.tohing period. o
The difference in the n\Dilber of days to hatching for eaoh raoe and
eaoh lot ns tested for signifioanoe by student 1 s ! . For Lots 1, 3, 4
and 6 (see Table 6 tor lot temperatures) the probability of the ! values
was leas than .01, that is, the differences in mean hatohing times be•
tween raoas at the same temperature were highly si~nifioant. The non-
nor.mal diatrlbution of the number of ~rs to h&tohing makea use of the t
test questionable and prohibita its use for Lot 2 data.
When the four r&oes are arranged in the order of the number of days
to hatching, the order remains the ~a at aver~ge temperatures of 39o0°,
50.60, 66.2° and 61.3~. Lot 2 data were not included for reasons given
above. The order, beginning with the raoe with the shortest time to
h::.tohing, is Sacramento, Green, Skagit and Entia.t. These data e.ra plot-
tad in Figure 13. The probability of these values randomly aligning in
1 1 l this order is (TJ)(l',H'!'!), o:r one ohanoe in S456.
For the conditions of Experiment III, the differences in the rate
or developnent between rao.aa are evident from Fie;ures 13 and 14 even
though aome of the ditferanoes are not great. With larger samples fran
each raoe, that 1 s, more spawning pairs, results different from those
obts.ined here would be possible if it so happened that the ·salmon in
these experiments were atypical representativee of their raoe.
In oonolusion, the incubation rate of the Sao~ento eggs in these
sxperiments was about 8 per oent faster than the Entiat eggaJ this
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
72
difference 1~ lignifice.nt. The rate of develo~ent of the Green and Ska-
git eggs was intermediate to the Sacramento and Entiat 'ggs.
(b) Threshold temperatureo !he threshold temperature and confidence
11mi ta were oaloulated for the four raoes by the methods given on page 60
and are tabulated in Table 13. Although the same order for the r&oes
that prevailed for the rates of development is present for the threshold
temperatures and confidence interval. the oonfidenoe intervals overlap
widely and lin1it the signifiaAnoe that may be attached to the ordering
effect.
0 0 The range of threshold tamperatl~es, 3lol to 32.6 F, for lots
reared at changing tsmperaturea is less than the value of S3.8°F for the
threshold temperature of lots reared a.t constant temperatures in the
range from 39.8° to 57.aor. To investigate further the difference in
threshold temper~tures between lots reared at oonstant temperatures and
at changing temperatures. the threshold temperature and confidence inter-
va.l were calculated for lots reared at constant temperatures at all
tompar~ture levels. In Table 13 the results of these calculations show
that the threshold tsmperatures of the lote reared at constant tsmpera-
tures are higher and lie outside the range of lots reared at changing
temperatures, but sinoe the confidence intervals overlap. the differences
may not be Bignitioant.
In conclusion, the threahold temperatures of the four stocks reared
at changing temperatures range from !1.1° to 32.60,. The range of c001-
pcr&ble values for egge tram the Green River stock at constant tempera-
tures is higher, !2.1° to ~4.0Gr, but the oonf1denoe intervals far the
two groups overlap.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
13
TABLE 13
E.atimatecl Threabold Temperatures and Ccn!idence Limits
for Xxperianta I, II and III
litiili&tid
hperi-Threahold
wt Raoe Temi?!rature* Confidence Limits
ni Sac ram~n to 32.6°1 3l.2-33.9°F
It Green 3l."f>r 0 29.8-33.3 F
II Skagit 31.6°1 0 29.8-33.2 F
" Entiat Jl.l°F 28.9-33.0°F
I & II Green, 0 33.8°F 0 33 • o-3J+. 6 F
Temp.,40-58 F
I Green, 32.7°F 0 30.4-34.5 F
All temps
II Green, 34.0°1 0 31.8-)5.8 F
m temps
*'I'hreshold temperature = i-k(~) where i ie tile re~ression coefficient
of the rate of development on temperature.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
74
( o) 1'empezaature SUDD&tion oonatant, k. Having dertermined the thresh-
old temperature, !.• the temperature s~tion oonetant, ~· was estimated.
'l'wo eatimatea were madeJ one in which the threahold temperatures were
the oaloulated values as detend.ned in Table 13, and a seoond in whioh
the threahold teperature na arbitrarily taken aa 32°F.
The ftluea of the tanperature constant are given in Table 14. When
tt&l.' oaloulated values of the threshold temperature are uaed it is seen
that the ~ values vary more b.tween races but have a smaller standard
error than when the thre~old. tempere.ture ot S2°F ia used. For eggs
fran the Green River stook the average Talue of the temperature s'I.IDmation
OOD,atant. when !2°F is the threshold tanperature, is 932, which is equi-
valent to 932 temperature units as defined by W..llioh (page 43). This
ia similar to his estimate of 900 temperature units to h&tohing for Sao-
r~ento River chinook aalmDn especially if allawance is made for the
more rapid rate of development of the Sacramento fish (page 72).
In oonolusion, the best estimate or the temperature summation eon-
stant. ~· is made when the threshold temperature is calculated f'rcm the
equation, a • 'i-k(!), where l/Jc 1& the regression coefficient of the y
rate ot deTelopment on temperature. However, if a is unknown, 32°F is a -
reuonable estimate of a. Using the oaloulated estimates of !.• the val-
uea of the temperature a~tion oonst&Jrt up to the time of hatching for
the eggs of Experiment III tram the Sacramento, Green, Skagit and Entiat
Riv.r obinook almon were 860. i60, 960 and 1020, reapeot1Tely.
(d) Incubation time of Experiment I aublota. Experiment I waa bas-
ioally a oonstant tcperatlure experiment. However, there were five
aublots of 100 egga each in whioh the egga were maTed during the inoubation
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
15
TABLE 14
Temperature Sunmation Constant, k
(y)(x-a) = i<
Range
~ri-No. of An. 0 m~t Stock Lots Temp. F &il' k-s.e. a k-s.e.
I Gr•n ll 40-65 32.7 919_ 5.8 32 960_ 9.1
II It 10 45-62 34.0 815_ 5. 7 II 897.Ju. 3
I & II II 8 40-58 33.8 828....10.2 11 939_17.7
III II 5 39-56 31.7 944-9.4 II 922-12.1
II Sac rWDIIIn to 6 39-62 3~.6 85.5-15.4 II 887-14.1
" Skagit 5 3'1-61 31.6 951-12.9 II 921-17 .o
II Entiat 5 39-61 31.1 1020-17.1 II 953-26.7
Average for Green River 877-3.97 <)30-b. 64
Average for Exper.iment III 941-6.ll 921-8.01
*Threshold temperatures from Table 10
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
16
period .fran medium to high water temperatures and three eublots in whiol
the move was .from medium to low water temperatures (see Table 15). Al-
though the eg;gs wore transferred for the purpose of observing the effeo·
upon moristio characters and upon the relation of relative change in
temperature to per oent mortality, the results also show an effect upon
the rate of development.
The rato of development at the average temparature of incubation
WilLa slower for the ::ublots moved to the nigh temperatures than for lots
reared at the oorreaponding constant temperature. For sublots moved to
the low temperatures there were no lots rear8d at the corresponding oon·
1tant temperature for oomparism, but the rate of developme:rrt of the
sublots waa as faat or possibly faster than expected from the projeotiol
of the rate of development ourve (Fig. 16).
All experiJilenta oanb1ned. An adequate expression of the rate of
development for lots reared at constant temperatures in the optim\.111 ran1
and for lots of the four stocks reared at ohanging temperatures has beel
found. In an eff~ to find a general empirical equation to fit all th~
data. even though several complexities m&y have been introduced by oan-
bining lots--irrespective of raoe, temperature pattern, year or mortal-
~ .... I ity--~he temperature summation rule. the Belehradek equation, and the
logistic curve were fitted to the temperature-development relationship
tor fifty lots from the three experiments. This inoluded all lots exce]
the two that were incubated in water ot low oxygen oontent. The values
for average tamperaturss and the number of days to hatching for these
lote are to be found in Tables 9• ll and 15.
The fit of the temperature summation rule in the reciprocal form t~
these data was tried b.Y plotting the relationship of the avsrage
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Lot
8C9
8Cl0
8C7
3A7
2A7
8Cl
8Ba2
8Bal
17
TABLE J.5
Water Temperature, Incubation Period am Par Cent Hatch of
Experiment I Sublets
Temperature History
At city water tempdrature
55 to 48°F, for 25 days
then to constant water
tem.t-erature of 65°F
Same as 8C9 except to
62~ °F water
Same as 8C9 except to
tiJ°F water
28 da~s at 45°F, then
to &J F
28 days at 40°F, then
to bfPr'
25 days ot city water
te.'!1perature, then to
34°F
ld days at city water
temyerature, th<;.ri to
34°F
.lJ. uays at city water
temperature, tht; n to
34°F
55.9
55.7
51.2
49.8
37.1
36.5
36.1
Days to
50% Hatch
40.3
4J.4
4.3.3
5li. 3
57.8
144.0
158.0
172.0
Per Cent
Mortality
93
28
3
3
2.3
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
.c:
(.)
..p
~
~
"U"\
0 ..p
10 ~
:::l
~
H
CD
!
8
r-f
~. 0 t8
J.So''-
).CC-
2.50-
2o00-&u .
• 6.
f .
1.50-
0
0
1.00-
.
~
• 0
"
.5o-
.OO·ib' -==
35 45 5o
Temperature, ~
0
0
0 0
0 0
0
o· l:. .
I' C·
~ .__.
o Experiment I and II
:~ Exp. I Sublets
Experiment III
'_,__~~L-_~0~
55 60
Figure 1). Rate of" I.Bvelopment and Temperature for All Lots,
All Experiments
0
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
79
temperature to the reoiprooal or the number of days to hatching (Fig. 15).
The relationship was not linear and therefore a ourve that fitted the
data more oloaely was sought.
The fit of the Bilehradek equation to these data was tested by plot-
ting the relationship of the logarithm of the average temperature to the
logarithm of the number of days to hatching (Fig. lSA). This relation-
ship also deviated from linearity.
(a) Logistic ourve. For the logistic ourve tho fit to these data
was tested by plotting the relationship of temperature to the logarithm
of L-P/P (page 42). As a trie.l run, the relationship of temperature to
the log of L-P/P were plotted for eight lots that were approximately
equally spaced throughout the t"lllperature range at which chinook salmon
eggs develop (Fig. l6B). Sinoe this relationship was practically linear,
the logietio ourve was then fitted to the data for &11 lots fram Experi-
menta I, II and III (Fig. 17).
The constants for th• logistic curve w~re determined by the methods
described on pages 42 and 6S. For L, values of P at 3°, 9° and 15°C
were used. The calculated equation for the maber of days to 50 per
oent hatch is
1~2.SOO-. 2022x
y = ----~~~----.M~04
For this equation the standard error of estimate, Sy•X' whioh is an esti-
mate of the t1 t of the calculated ourve to the observed data, is 3 .14.
The reoiprooal of Z• t~es 100. is ths per oent development per day for
which the equation is
100 -4.404
Y _1 +;2.!6~-.2o22X
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
2.20 IL
e:,
(f)
~ < &:! 2.00 .~
~X.
0
16 ..... E-t
~ ~1.80 .~
E-<f;
~~
~[5
~p..
~gl.6o §~
'I :I
I;
80
e:,
e:,
0
' .
0
0
Bel.ehradek
.
Ooc. .
A
l.ho lb----c--~~-
1.55 1.60 1.65 1.70
L(X} OF AVERAGE TEMPERATURE, °F
B
Logistic
0.6!-
X
-1-~~
I~
"
I
i.4l~~------
4o 45 50 55
X
60
Figure 16.
AVERAGE TEMPERA TtiRE, °F
Relati<nship at the Beleb:rS.dek Equation and the Logistic Equation
to the Rate of Devel.opment Data from all Lots o A, Relationship
r.>f the Logarithm of Temperature to the Loga...'T"ith:m of the Number
~ Days to Ha.teh:l.ng. B, Relationship o£ Temp€rature to the
Logarithm o:f L-P/P.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
81
175 1 ~
/
13.5
l+e2.Jo6-.2022x
'I
I
I
.cihd.
l50;~ 100-1JoO
/ y :
4.4o4
~ 125~ l-re2.306-o2022x ;
ll r, --<!2.5
~
0
E-t
n!
~ 1ool l ;2.0
ffi p.,
0
l..C\
0
8
~ ~
~
I
l
i
l
·' j
75t il.5
1
50~ jl.O I
'
1
d
l
1 25 ~ 0.5
o~~----~------~-----~~----~-----~------~~0 JS 4o 45 So 55 6o 65 o
'l'EMFERATURE, or
Figure 17. The Relat:icmsh:lp a£ Temperature aDd the Number of Days
to Bate.h1Dg to the Logist.ic Curve and It.s Reeiprocal,
for All Lots
§~
" ~ ~
~
I
§
~
0
~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
82
In conclusion, when data traa all lots in the three experiments are
uaed, the equation of the logistic curve in the f'orm
l+e2.306-.2022x
y • .04401 '
where z is the number of daya to hatching at temperature ~' best describes
the temperature-denlopment relationahip. The standard error of estimate
far this curve is S.l4.
(b) Dut"ati~n of hatching period. The duration of the hatching per-
iod ns m.asured by the number ot days between the hatching of the fifth
percentile egg and the ninety-fifth percentile egg and was called the 5
to 95 percentile range for the hatching period or chinook salmon. By
not using the f'irat and last five per cent at the total ranr,e, the few
very early or late hatohiili.; eggs that oooasionally would ooour were not
included. The relationship of the temperature at tirne or hatching to
the 5 to 95 percentile ran~e for the hatching period is shown in Figure 18.
The duration of the hatching period might be expected to be influ-
enced by the r&te of devela~ent and thus to decline w1. th increase in
temperature, but this was not exactly true as shown in Figure 18. From
S5° to 40<7 the duration of the hatching period rapidly declined, but
above 40° the length of the hatching period was short and without notioe-
able change with respect to temperature. The range ot average tempera-
~•• for which the mortality of eggs and fry of the chinook salmon is a
minimum, is also the range for whioh the duration of the hatching period
is a minimum. It would appear that & short hatohing period ia associated
w1. th a high survival.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
A
e;.
Jo;~
(f)
>-1 ~ 00
~ 25-
r:i:l
0 0
~ 20-0
0 0
~ u 15-E-1
;:j 0
t9 0 H 10-~ 0
u 0
~ 8 0 0 ~ 5-0
0 8 0 0 0 0
1J'\ 00
00 0 0.. 0 0 0 0 ~~ ~ 0 • 0 0 0 0 1J'\ 8 0
o-~----J .. ~------'---_
35 40 45 so 55 6o 65
TDfPERATURE A'! TIME OF HA.TCIITNG, OF
Figure 18. The S to 95 Percentile Range for the Hatching Period of
Chinook Salmon
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
84
Yortali tias
=
Lots reared at constant tem_Feraturea. The weekly cumulative mortal-
1 ties of the lots reared at constant tenperatures in Experiments I and
II are shown in FigUl'e 19.. For lots reared at corresponding temperatures
in both experimenta--60° ,66°, 50°, 460J'--the graphs up to the tenth week
' (the end of Experiment II) are &Terages of the two experiments. By in-
apeotion of Figure 19 the lots oan be claasified into four groupe as
t'ollawec
1. Lots in which mortality during incubation is 100 per oent, that
is, no hatch. This includes the lots re&red at average temp(ltratures of
2. Lots in whioh a fsw survive to hatohin~ but die in the yolk-sao
st&gdo In this category are the 62.6° and 60°F lots.
z. Lots in which the mortality to hatching is low but is followed
by a high mortality during absorption of the yolk sao. After feeding
has begun, mortality is again low. The 57.6° and 660f lots a.re in this
group.
4. Lots BUoh a a 60°, 46° and 40°F • in whieh mortallty is low during
inoub&tion, yolk-sao and fingerling .rtap;es. This is the optimum temper-
ature range in reapeot to mortality for ohinook sa~on reared at constant
tGperatures.
One explanation of the high mortality that occurred during the ynlk-
sao stage to th~ lots reared at 67.5° and 55°F is that the or~nization
of the physioloEioal prooeasea ie out of step. Hayes (1949) wrote that
phyaiologioal processes have optim\Jn tnnperaturee whioh vary with the
prooeaao For example. bile formation is fa~ored at high temperatures
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.l.OO · 174° 671° f1° .............. 621° -~ _-6o0 I I ••••• / . • i •• / • I : I ' ( ~~· I I : / I : / , I : I ) 5 ----0> I : 80 . I i I . I I I t:. ,' i ( fiil • t : I 0 I I : I ~ I I !i: . : : I ~ 60. I I i I ~ i f 1 I ~ j / ~ I 1 end of ExperiMnt II & ~ 4o· j ! j I / 1 ,. : i I s71• I . I :I I .,-40 I : .·· I • / I I .. t) I I ./ 20. ) I i I ... -... ----·· I / i · I r-: so• . , . I \ I_~/ 1· I ___ __ _ '--us• ~,t~ 1/ ~ ~ ..... ..;--...-·-~v .. J_ ---· •• -···-··..-, ,• ••••••• .JI"/-. ----... -···-· 01'............... --_ ... -... -( ·~~----··· ··· ------------· -----·---1 r ·--------·-•· ---··----0 ' 10 15 42 46 WEEKS AFT I!R START OF EI:Pl~UMDrr Figure 19. CUIJIIlla.tivo Mortality in Par Cent by Weeks tor Lots Reared at Constant Tempero.tures in Experiments I and II
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
86
and circulation in the yolk ~t law temperatures. Thus, exposure of the
egg to &n unfavorable tenper~ture may not result in death until the yolk-
A.O stage.
The relationship of tanperature to per oent mortality at time of
hatohing i1 shown in f'ig\U"e 20. To supplement these observations, espec-
ially at low t«nperatUJ"es, Burrowa' data for sir.:i lar experiments with
ohinook sa~on at the Entiat Hatchery are included (Table 16), The rapid
inorease in mortality at temperatures of 60°F and higher and at tempera-
tures lower than .f.OOJt' are to be noted.
From Figure 20 an approximation or the "lethal temperature 50 per
oent, u 50," wa1 made. This ia the temperature at whioh 60 per oent or
the individuals die from temperature effects. Taking into account the
morta.li ty not due to temperature, whioh wa.s ass1.1!Ud to be the average
mortality in the optimum range (4.6%). the LT 50 was the temperature at
the 55% mortaU ty level. The ourve of morta.li ty in Figure 20 crossed
the 55% level at two places, 36.5~ and 60oB°F. whioh are the estimates
or LTso•
Changing temperatures. The egg mortalities for the Sacramento,
Green, Skagit and Entiat races are listed in Table 11. From inspection
ot the table several facts are evident.
First, the high mortality of the lots fram the Green River atook
waa not due entirely to temperature. In Lot 5 the egg mortality was 100
per oentJ for the five other lots, the mortality was at least ten times
as great as the averap;e mortality for the other three raoes1 therefore
it is evident that same ot the mortalities to the Green River lots were
tram causes other than temperature. Also. at an average tanperature of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
~
!j
< ~
:;:E:
~
0
~ ~
87
lOQi!-0 " ><
0 E:r:perilllents I and II
Bo·r-X
Entiat Data (Burrows r) I
6o~
4o~
0
"
2Q;~ 0
"
0 " ><
..... 0 " >< 0 "o
Ol~. " )l ~ )( .o o, 0 . "' )<. .I
35 40 45 50 55 60
TEMPERATURE AT TIME OF HATCHING, ~
Flgnre 20. Temperature and Per Cent Mortality f'or Lots
Reared at CCIIlSt.ant Temperature
-
~ 0
0
0
~~
65
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
88
TJJ3LE 16
·r'iater Temperc:.ture, IncubaL un Period cnd .Mr.Jrtality o: C 11incok .::ial.aon Z15
Reared c.t Constant 1erep:~rat"..!res a.t the Entiat H~\t<.:ht.Xy (c,uro-ws' Data)
Year
1952-5.3
195.3-54
1953-54
1955-56
Days to Fer Cent
.. Brcc:>d .:>tock __ }~~·~0_F. ____ 5_0%_~a_tc_h_. ___ &~_o_r_t_al_l_·t_y_
Entiat
Entiat
Skagit
Entiat
49.78
54.38
57.53
59.61
35.10
37 . .35
40.05
L,2.. 64
44.89
)4.39
37.29
40.04
4.~. 54
44.87
39.94
42.40
44.74
47.38
49.21
52.19
41.88
)6.6'-}
J4.34
204.0U
1.57.54
l2G.11
';12.38
76.82
206.:.;3
160.29
u:.L49
94.00
76.32
1~ .10
94.69
78.91
63.40
5) .44
7.1
5.7
6.1
1~.4
99.6
52.6
l~~. 5
6.1
18.4
98.7
30.9
10.2
2.1
0.9
2.7
1.3
0.7
0.6
1.1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
89
620p, the higheat average temperature for lots of Experiment III, the
effect of temperature upon mortality is marked. From 66° to 62°F the
increase is about 20 per cent and corresponds to e. simi la.r increase at
aorreaponding temperatures for the lot• reared at oonstant temperatures.
Finally, there is no difference in the tolerance to law temperatures of
Sacramento, Skagit or Entiat stooks as shown by egg mortalities. The
minimum temperature was S40p and aozne lots from all races were incubated
at th.:ia temperature for twenty days, the descent and ascent fran the
minimum being one degree every five days.
These data are not adequate to define temperature tolerance. The
tolerance of ohinoolc almon eggs to limited expoeu.rea at high tempera-
tures has been investigated by Donaldson (1955). He f'owd that the
exposure time neoeaaary to oauae 10 per oant kill averaged li, 4 and 13
days at temperatures of 67°, 660 and 63or, respectively. For fingerling
chinook salmon Brett (1952) has determined the temperature tolerance.
Abnormalities
The term "abnormal fish" is difficult to define. In this report
the definition is limited to individuals with morphological abnormalities
that can be reoogni&ed visually. For the egg stage per cent mortality
is a good measure of abnormality, ainoe any egg that fails to hatch is
abnormal, strictly speaking.
For the fry--the stage fran hatching to feeding--mortalities were
olaaai.fied as to type by the terms used by Foster ( 1949) to identify
abnormalities in the progeey of rainbow trout exposed to X-rays. These
~s include the types of abnormalities that were found by Welander
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
90
( 1~) to ooour in the young of ohinoolc salll:Lon exposed to X-raya in the
egg atage. Identification of abnormalities was made frillll preserved spec-
imens.
In Table 17 the per oent and number ot different types of abnormal
fry in Experiment I are aUIIIID&l"hed. Far the few individuals with two
abnormalities. both typea were recorded. The number of abnormalities
increased at the extreme tenperatures. but were principally abnormalities
such as "developmentally deficient." "weak body structure" and "serous
fluid" rather than the monster-like abnormalities of "spinal curvature."
"distorted jaw" and "twinning."
Growth
Eggs from lots of ~periment I were weibhed before and after water-
hardening and near the mid-point of the incubation periodJ fry were
wttighed and measured onGe, just after hatohingJ and the fingerlings were
weighed at two...,eek intervals from Yay until October. Re1m:ts of the
measurements of the eggs e.nd fry are summarized in Table 18.
The rate of water absorption by the egg was measured by plaoing 30
egg1 in a ruled trough and observing the total length of the row at five-
minute intervals. Arter 35 minutes in water the eggs had reaohed maximum
size. At the time of placing the l;,ggs in the water the diameter was not
determined, as the egg a were sort and s01Jl8What irregular 1n shape. Ai'ter
absorption of water the eggs were firm and spherical and the average dia-
meter of a sample or 30 eggs was 9.S mm. The inor-ea.ee in wei~ht during
the water absorption period was 15.0 per oent as determined from a sample
o£ 136 eggs tha.t averaGed 379 mg before and 436 mg after water-hardening.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
71
TAblE 17
Abnormal Fry In Let. s Reared at Conslc;nt Ten.peratures
======= ==--c-.-:c--· _-_ -==--~-=,.---·-·----
Lot
Nu..'llber
Hatched
lt:r Cent
Abnormal
2
40
368
8
3
45
352
Number of hbncr~alities by Type*
D 5 J
fl 16 3
s 3 l
c 5 0
T .... l 0
J 0 0
E 1 1
T 0 0
4
City
Water
501
6
lO
7
2
5
-~
_.)
1 -
.L.
.:.
5 6
50 55
440 480
2 5
1 .2
0 ~
1 0
u C3
(l 4
G G
(j G
J 6
7 10
bO
404 120
40 65
53 32
45 10
32 JJ
19 ()
11 6
G u
3 5
1 0
* D, developmentally deficient; "', we<.ri: bcJy structw·e _; S, serous fluid; C,
SJ.:;inal curvature; L, shortened bu~; J, di:'ltorted ja.w; h, defective eye;
T, t wi.'1Il :_ng
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.Table 18 Average Weights and Lengths of lgga and Fry From &x.f,!eriment I Weights in mi.lligram.a and 1angtha in mill:imetera of !o:rmal.in preMned •v.aimena ---~---------------------------~------· --------------------------~---------· ----------------· --------------------~----Lot Temp. n Total VJei.ght Yolk Wei6bt Shsll \'iei#It Fry W.ight Fork Weight ------------------------·-----------------~----.. ---~ 2 40 10 4ll ~ 4.9 JlJ .: 4.0 Zl.6:!:. 1.14 approximate midpoint 3 45 a> 412 ±. ).2 319 + 2.2 Z/.0-+ .bO 331 ~ 3. 2 -or 4 47* 20 402:!:. 3.4 26.0 ~ .44 hatching 5 50 20 410 :!:. 3.5 JJ5 + 3.4 22.2 ;t .57 6 55 20 402 ~ ).2 332 ~ 2.6 24.1 + .Zl -7 60 20 403 :! 4.1 344 :. 3. J 2J.4 :i .)3 '0 8 47* 20 404 ;!;. ).8 ))2 1. 2. 9 18.4-:: • 55 N 10 62 2) 411 ± 3.4 355 !. 1.8 18.3 ~ .35 9 65 20 398 ±. 2.8 351:!:. 2.5 16.4 l. .32 for &l.l lots 170 405 !. 1.3 J36 ~ 1.4 21.7 :t -~ ~ 2.60-+-2.9 juat a.tter 2 40 10 346 .± 2o) ao.o :::. 4.2 24.0 z .40 hatching 3 45 10 .340 j; 1.0 276 ;i ~-4 58.5 ;t 1.9 22.3 ~ .13 4 47* a; 334 .t 1.8 268 .::. 1. 5 63.3 -+ 0.9 22.9 !. .12 5 50 20 321 ::! 2.5 266 :t L9 49.6 ~ 0.9 2.1..8 +-.23 6 55 20 337 ::!;. 2.8 2?5 :!: 2. 7 59.2 :i o. 9 22.0 ~ .08 -for all lots 80 334 ::!;. 1.1 270 :!;: l.U 59.8 j: 1.1 22.5 :t .08 *Average of city water temperatures
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
93
For the samples withdrawn fram the egg lots at the estimated mid-
point of incubation the weight of the yolk as listed in Table 18 also
inoludee the weight of the embryo. These values are not oomparable frCI!l
lot to lot beoause of two unoorreoted errorsJ one is the error in esti-
mating the mid-point of hatching, the other is the loss of weight due to
dehydration of the egg during the weighing prooeas. The error in esti-
mating the mid-point of hatohing ranged .from minus 28 pGr oent to plus
16 per oent and was determined by eubtraoting the number of days to the
mid-point of hatohing from the number of days to the time when the sample
waa withdrawn, and dividing this value by the nlDlber of days to the mid-
point of hatching. The lose of weight tram dehydration for an egg before
removal of the shell was at the rate of 18 mg per hour. While waiting
to be weighed the eggs were subjected to dehydration for a p~riod cf a
f6W minutes to one-half an hour, a loss in weight of perhaps 1 to 10 mg.
However, the inverse relationship between tamperature and shell weight
11 probably true even if consideration is given to these two errors.
The weights and langths of the newly hatched fry deorease with tem-
perature but in an irregular manner. The try tram the 40°F lot were
definitely larger thAn the try from the lots reared at higher tempera-
tures, whioh agrees, in general, with Gray' a observation. For Salmo
tario Gray ( 1928) made the following statement, "When eggs are inoubated
at low temperatures the anbryos at the moment of h&tohing a.re signifi-
cant~ larger than those hatohing trom eggs inouba.ted at higher tempera-
tures.• This is also probably true for the ohinook sa~on, but it is to
be remembered that the !'ish reared at lower temperatures are also older
at the time of h.a.tohing, the age for the 40°F lot being 128 days as
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
94
oaupared to 50 days for the 60°F loto
During the fingerling stage lots were weighed and oounted at approx-
imate1y two-week intervals. Five lots survived to the fingerling stage,
inoluding the control lots whioh were reared at oity water temperature.
The fish were weighed as lots ra~her than individuals, as it is not feas-
ible to weigh'live fish of this size individually.
To investigate growth rates of fingerlings at oonstant temperatures
above 65~ a oontrol lot was subdivided on May 1, five weeks after the
start of feeding, into four groupe of 100 eaoh. one group was retained
at oity water temperature and the other three were transferred to water
0 0 0..,. temperatures of 6U , 67 and 74-r. Temperatureii were raised at the ra~
ot one degree per d.a.y from the oity water temperature of 54t>r' on l4ay 1
to the temperature selected far the lot.
Growth curves for the original lots and !'or the lots started on
May 1 are •hawn in ligure 21. The difference between lots ia obvious.
From 40°F upward to 550f the growth rates increase and from 6U°F upward
the growth rates decrease. The average we1Eht at 46 weeks for the 40°,
45°, 50° and 56°F lots was 0.4, ~.~. 12.6 and 18.1 grams, respectively.
For the lots started May l the average weights were 11.2 and 7.5 grams
tor the 60° and 67°¥ lota. The 74or lot did not surviveo
The maximum growth rate for .fingerlings rl!lared at constant tampera-
turea ooours at about 55°F, but the fastest growth rate shown in Figure
21 ia tor the lot reared at o1ty water temperatures. This was observed
during the 3oth to 32nd week at water temperatures of 60° to 63°1-', but
tince it is reasonable to assume that there is a short lag in the re-
aponse of growth to temperature, the optimum temperature l'or this lot is
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
95 city
.... -·--water
Previous to May 1, all at
City Water Temperature
:IS-.·
.·
Q .c __ _
0
' oi
0
' . "-·-~---•OHo• .1..-----· 0 , oO 0 -~ ••
,_ -'
15 20 30 35 40 45
NUMBER OF WEEKS AFTER START OF EI.PERIMEN1'
49 54 60 66 64 64
l J l L __ -_J
city water temperatures, Op
5SO
Orl.gj naJ Lots
-~-: /.-·-city
•••• l_""'
5
/·/ water
/·_ ... ·· / ........
.--. 50° // /
/ ; ....... /
/" ..... /
/ _.....-/
/' ....... /
. / ... ~
/ -. ...----·4SO
. /, --./ .. ·· ----___..._.... ---::.....---: ... ··········------40° . . . __ ......_ ____________ _
_ __.....___ ·----1-..... • I 1 • l, •• M ------·'·-
20 25 30 35 40 45
NUMBER OF WEEKS .AF.rffi START OF EXPEruMENT
Figu.:re 21. Vaight Curves far ExperiJDent I
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
96
likely to be during the 28th to 30th week at temperatures of 57° to 60°F.
The effeot at mortality upon grawth rate was !light for the lots
that survivad fran the beginning of the experiments (November 15) • e.s
fingerling mortality was less than 6% except for Lot 2 which failed to
feed. In the experiment started May 1 the 740f lot died in 15 weeks and
the mortality was greater than ao< in 22 weeks for the lot at 67or.• The
growth rat~s for the 74° and 67°F lots are less reliable for this reason.
In conclusion, the opti~um temperature for fingerling growth is
between 55° and 60°F • The growth rates by temperature lots decrease in
regular order on either side of the optimum.
Meristic Characters
The claaaifioation of fishes especially as to species depends to a
great extent upon the count of meristic characters. Also, the use of
meri•tic charaotera. particularly vertebrae, became a widely aooepted
method for defining raoes after Reincke's investigations in 1898 on the
raoea of herring.
Even before Heinoke•s investigations the ~eographioal differences
in vertebrae number within species had been associated with temperature.
Gabrial (1944) wrot& as follDWlla
Following the early ~eneralizations of GUnther (1862)
&l'1.d Gill ( 1883) that the number o!' vertebrae is
higher in genera of fishes inhabiting northern lati-
tudes than iu relsteu fiehos from tropical re~ions,
Jordan ( 1891) prepared a •law" setting forth an in-
v~ae relationship between the vertebrae number o:f' a
species and the water temperature prevailing in ita
geographic ra~e.
~he upper lethal temperature limit for chinook fingerlings aa stated by
Brett (1952) is between 24 and 24.5°C (75°-76or).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
97
From Jordan• a "law" and Reincke's work the idea developed that raoial
differenoes could result from the effeot of temperature upon meristic
oh&ra.oters.
A great many racial studies oonfinn Jordan• s "law." A ffJW of these
are HUbbs (1925), Rounsefell and Dahlgren (1932), Tester (19S8) and
McHugh (1954b) on the herringa Schmidt (1930) on the coda Weisel (1955)
on the oyprinidsa &nd Mottley (19!7) on the trout. How8Ver, racial stud-
ies haw two shortcomings when used ·:.o demonstrate the effect of tempera-
ture upon meria"tio oha.raoters. One is that temperatures durin~ development
are estimated, not known, and secondly, the oounts of the meristic ohar-
a.oters of the parents tl.l"e unkncnm.
Laboratory experiments on meristic characters of fi~h are few. Tin-
ing (1952) reviewed these experiments, which inolude Sohmidt (1917, 1919,
1920 and 1921), Mottley (l9S4 and 19S7), Tining (1944, 1946 and 1950),
Gabriel (1944), Heuts (1947 and 1949) and D&nneTig (1950). To this list
Marolcmann (1964) and Lindsey (1964) should be added. Conclusions frcrn
these experiments are that either low oxygen or high COg pressure increases
the number of vertebrae, pH in the range 6.4 to 7.8, egg si&e, fry size,
or early or late hatohing have no ef'feot on vertebrae numbers and salin-
ity and temperature modii'y both vertebrae and f'in ray number.
Modifioation of vertebrae number by temperature has not been oonsis-
tent in tM la.boratory experiments. Schmidt ( 1921), Tining ( 1950) and
Lindsey (1954) have shown that the lowest number of vertebrae ooours e.t
intermediate tanperatures while the results of expnriments by Gabriel
( 1944) and Dannevig ( 1950) shaw an increase in manber of vertebrae with
deoreaeing temperature.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
98
In the pr a sent ex per i.meot a o ount s were made or vertebrae. dorsal
rays and anal rays to investigate the variability associated with tamper-
ature that oooura in the meristic oharaotera of the chinook salmon.
Vertebrae. The number ot vertebrae was determined by counting th11
oentra between the baaioooipital and the urostyle.• The vertebrae as
seen in a radiograph are shown in Figure 22. When abnormal vertebrae
were encountered. the number was determined by counting the arch elements,
but when both the oentra and arch elements wel"e in doubt, no cowt was
made. In the caudal al"ea. the centrum was counted as one if separation
was not complete.
The vertebrae counts of lots from Experiments I and II are reoorded
1n Table 19 and shown graphically in Figure 23. The u-ah.aped curve of
Experiment I shows that the number of vertebrae increases at both high
and low temperatures and is similar to the findings or Sohmidt (1921)
and Tining (1952) tor the sea trout and Lindsey (1954) for the paradise
tiah. The data for Experiment II are limited to the high temperatures
but substantiate the Experiment I data for those temperatures.
For :&xperiment III the reoord of vertebrae counts is tabulated in
Table 20 and is shown graphi0&11y in Figure 24. These data do not shaw
the aame inorea.se in the number of vertebrae at high and low tsmperatures
as was seen in the Experiment I dat&J on the other hand, there is no de-
crease in vertebrae number at high temperatures auoh as was found by
Gabriel (1944) and Dannsvig (1950).
The temperatures gi~n in Table 20 and Figure 24 are the average
values during the incubation period but moriJ properly should represent
•Vladylcov {1954} ata.tesa "Uroa?'le is the posterior terminal segment
whiah tollow.:J the last undou'bte centrum. In Salmon1dae the uroatyle
remains non-ossified."
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.~ :0 ........ • .............. .. ............ -• 30 ~'-, ·. ' ' "-'-'f' ' '\ , , 10 \ ·, I -JU1~a·u·n~~· -· · 1ltllr: ... ·-.......... -······ · ·-· -~-. . ; . ". , ...... :•"" ,.,,..,,.,.,. .. ;., .. (:;;,~;;~·;"'·"""~ "" . '/.;~>/;( . . ; . .,:c:<i-~:, /~''*;.~ 6 . 'P vertet.r::..e, 7) \ j,?. c.:.bdcn~in<-<1 + 41 caudd.l J anal ray::;, l') dor::>al rc..ys, lc; Figure 22. nadioe:;rc.1 h ui' a Ch:i.J1co~ J<.tlmc)r. Finr;erling ( x 3~) ------~c 'The first caudal vertebra of a salmon is defined as the verteLro. with a "sudden increase in leOF~th oi' the haemal spine •11 The r irst haernal Sf;ine is indeterrninCJ.te i:l s it j_s a :ninu te }JI'Ocess which r;radually 'uecomes lon~er on succeeding vertebrae (Clotr1ier, l';t50). '-!) ~
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.T.i.~LE 19 Vertebra.':.. Cou!1ts ..:-~· ~reen niver Chincoi( Sa:..:nun hec.r~.:d at Constant anr:1 City Watt:r Te·np:o:r<.ltures 'Ie;:liJ., NtL'I!ber of Vertebrae s Ex.p. Lot oF 66 67 6B (Jj '/G 71 X 1l X. l 2 )' J. F: ·, ·~ .:._, 135 59 1 69.17 2}8 .CJ410 3 I ' '7 , 109 84 2 68.44 196 .0377 --+4·. .l ) 5t, .I) ; 175 79 6R.30 256 .0298 () 5::-.. 1 , 59 44 6P..4l 104 .0504 j_ 7 60.~ l 2 6 3 1 69.08 13 .288 1.. IJ,7. ·~;; ? 1·-;~6 75 t..8. ;24 278 . CJ...-:91 8 1.. 7. C* ') '2.49 103 1 68.25 JC5 .U268 -~ II J 54. f_) 'l ';0 ')') b ::,7. 1+4 175 .0476 j..J .-8 5 5'' ' ~~ 63 117 18 67.7b 200 .0440 ' . ' '• 5~3. 5* )_ '!l 93 :22 07. 7'2. 188 .0495 -r, City Wd.ter
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
I
~
0
&i
i
68.5"
68.0"
67.5 c
Q
\
\
\
101
' \
\
----Green
xX
o constant temperatures
x city water
---· -=-co· -l,"'-. • -•--~··._,-, __ .,._,,.
40
-~
-.o···
50
/
.---d
~tr
/
I
/
/
I
I
I
/
/
I
p
.~II
I
I
d
55 6o
Figu.1"e 23. J.-nnge 1lamber of Vertebras and 'f8ilq)8rature £err
Lat.e t4 Bxperime:rit.e r and n
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.TABlE 20 Vertebrli:. Cuunt-:3 cf Jacr~nto, Jkae;; it, Green and ~ntio.L River Chinook So.lrnon uf Experimtnt III Lot Temp., °F 63 64 65 66 67 Sa 1 39 .o 5 :1.6 44 18 2 41.6 1 7 51 67 3(.1 4 56.5 1 23 ll7 98 5 62.2 ..... ll 14 j 6 55 .5* 16 108 133 36 Sk 4 55.9 1 25 5 ~;l.J J 20 6** 'J4. T:. ~ .) ~ l )9 • I~) J 2 '"l. 6 2 1 4 56,J 2 6 55.!t• l 1 } E 1 38.8 3 43.4 4 55. 2 5 t,l, J 6->:--il-55.2 OF' * At cc,nstant temperature, 54-56 Number of Vertebrae 68 G9 70 71 1 ,g 6 3 13 1 1 3 1C4 dl 13 JO 15 3 6 18 )3 '....4 4 26 46 ~3 5 9 4 12 10 c; ./ 14 40 25 2 10 39 5 1~ 3 ~1 5 65 2 8 ** Water of low oxygen content duril1t:, lncubatkn -72 TJ 74 75 76 77 ?8 X n 65.93 84 66.09 182 66.39 2.52 66.35 34 65.67 296 68.36 '2.2h /..;7.93 71 15 70.09 101 69 .8<) 99 69.43 21 1 69.57 30 69. C/1 84 :!.9 4 7l. 3J 82 6 7l.v3 JO 2:..7 29 3 l 71.73 4JL_ ll4 56 L. 2 3 1 7 .... 1... .. !.57 13 Ll 107 137 70 28 1 74.72 392 8 -X .091 .089 .047 .163 .046 .052 .109 .130 ...... .081 2 .320 .233 .104 , .... ,.., '"': • J'~J .355 .031 .U64 .c66
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
i ~
~
i
72 ~
II ~
71 t
~
'I
II
,I
10 l
:t
I
~ l .,
!I
'I
:i
" 69 !~
ll
,.
68f
~
II
'I
I
I
i
67 i
I
66t
II
:i
I
65 I
103
"""'------v-··-
-··-··Sacramento
--Entiat
----Green
----Skagit
------.... ----... .___. ~-----
o-··
.-----o
o..._
" -----"""0
_ .. --o----·--·--o ---
o average of changing temperatures
x well water
--*=-·=
40 45 5o 55 6o
Figure 2h. Average Hamber of Vertebrae and Temperatures for
!Dts ~ El:periment m
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
104
the temperatures at the time the number of vertebrae is datennined. Till-
ing (1952) has ehown that the plastic period for determination of the
number ot vertebrae in Sa~o trutta trutta ie from 146 to 165 D0 (day
degrees with temperature in degrees centigrade) for which the total inou~
bation period is 400 Do. Using temperature values calculated on the
balis that the plastic perioci for chinook eal.lnon is the same as for Salmo
trutta t:rutta,. the curves exprealling the relationship between tempera-
ture and the number of vertebrae in Experiment III were shifted to the
right but changed only slightly in shape.
To provide information fraR whioh the plllstio period for vertebrae
formation in the oh1noolc salmon oould be established the eight sub lots
or Experiment I were transforred to water of either higher or lower tsn~
perature& at ve.rloua times during; embryologioal develol!Jlent, Fry fran
same of these lots survived to & ai&e suitable for staining or radio-
graphing but were too f~ to make accurate observations oonoerning the
plastic period for vertebrae formation. However, in Experiment III the
data fram Lot E 6 suggest that the plastic period begins before the 21st
day for eggs that hatoh in 46 daya. Lot E 6 wa• incubated in water of
low oxygen content for the first 21 days, after whloh the oxygen level
was nonnal. The number of vertebrae in this lot was greater by 2.6 thar.
in any other Entiat lot at either higher or lower temperatures, and for
th1a reaaon it is believed that the plastic period for the vertebrae of
the ohlnook ealmon of this lot began before the 2lat day of inoubation.
From the hperiment III data in Table 20 the great n.riabill ty in
number of' vertebrae between stocks oan be seen. The racial averages of
the lot• for all temperatures are about 66 for S&oJ'111Jnento, 68 for Skagi-;,
69 for Green and 72 fo~ Entiat. By Ginsburg's (1938) definition the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
105
difference between the Sacramento and Entiat raoes is equivalent to a
species difference since the overlap in the vertebral counts of the two
races is lees than ten per oent.
The number of vertebrae was generally greater than the 66 reported
by Jordan and Evarmann ( 18 9t1) for the speoi es. From the r aport by Foer-
ster and Pritchard (1935) the average number of vertebrae for chinook
salmon was calculated to be 69.10 i 0.14, a.nd from Townsend ( 1944) the
average was 67.4. The range for Experiments I, II and III was 63 to 77
(for lots other thanE 6).
There was a marked increase in the number of vertebr&3 in Lots Sk 6
and E 6 which wer6 accidentally incubated in water of low oxygen content.
These lots were not included in the average values for the Experiment III
data because of the abnormal conditions. The average number of verte-
brae for Sk 6 was 1.7! greater than for any other Skagit lots and for
E 6 the inorease was 2.62 over other Entiat lots. Since there were no
other obvious differences between Lots E 6, Sk 6 and other lots, expo3ure
to water of low oxygen content is assumed to have caused the increase in
the number of vertebrae. There is subatantiatine evidence as to this
conclusion frOlll Tlining ( 1962). who .found that law oxygen content during
incubation increased the number of vertebrae. In the range from 58 to
98 per oent oxygen saturation the increase in vertebrae of the sea trout
was about 0.1 for each 10 per cent decrease in oxygen saturation (see
Fig. 7, op. oit.). The effect of higher or lower oxygen levels upon ---
number of vertebrae is not known.
Under the oonditiona that existed in Experiment III genotypio vari-
ation in the number of vertebrae was greater than phenotypio variation,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
105
difference between the Sacr~ento and Entiat races is equivalent to a
species difference aince the overlap in the vertebral counts of the two
races is leas than ten per oent.
The number of vertebrae was generally greater than the 66 reported
by Jordan and Evermann ( 1896) for the species. Fr·om the report by Foer-
ster and Pritchard (1935) the average number of vertebrae for chinook
salmon we.s calculated to be 69.10 l 0.14, a.nd from Townsend ( 1944) the
average wa.a 67.4. The range for E7.periments I, II and III was 63 to 77
(for lots other thanE 6).
There was a marked inorease in the number of vertebraa in Lots Sk 6
and E 6 which wer6 accidentally incubated in water of low oxygen content.
These lots were not included in the average values for the Experiment III
data because of the abnormal conditions. The average number of varte-
brae tor Sk 6 was 1.7! greater than for any other Skagit lots and for
E 6 the increase was 2.62 over other Entiat lots. Since there were no
other obvious differences between Lots E 6, Sk 6 and other lots. expo3ure
to water of low oxygen content is assumed to have caused the increase in
the number of vertebrae. There is substantiatine evidence as to this
conclusion from Tining ( 1962), who .found that law oxygen content during
incubation increased the number of vertebrae. In the range from 68 to
98 per oent oxygen saturation the increase in vertebrae of the sea trout
was about 0.1 for each 10 per cent decrease in oxygen saturation (see
Fig. 7. op. oit.). The effect of higher or lower oxygen levels upon --
number of vertebrae is not known.
Under the conditions that existed in Experiment III genotypic vari-
ation in the number of vertebrae was greater th&n phenotypio variation.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
5-
Green
o canst~· temp.
X city W:.e fl
107
X
X
... ___ ;__. -------·-------! -----
45 50
TmPERATURE, ~
-··-Sacramento
--Entiat
X.
60
20~-Green
15-
I
0 i -
----Skagit
o ave. of chang. temps
x veil water
p.
I ---. .· 'I
40 45
. / . / I / : /
....._ X I 1/
··~ . / .. i/
--· "'""6
..________
' ------<r-. --
/
/
50 55 60
0 ,
Figure 25. Temperature and Per Cent of Chinook Sa..lmon with Abnormal
Vertebrae far Lots of Experiment I, II and III
I
II
III
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
100
about one vortobr~ greater in the offspring than in the parents. If
this differenoe is duo to phenotypical variation oaused by some factor
in the environment. it is assumed that the effect ia equal for all lots.
In oonolusion. the laweat number of vertebrae are found at the
intermediate temperatures in the range fran 45° to 65°F. The average
nwnbeT of vertebne is about 66 for Sacramento. 68 for Skagit. 69 for
Green and 72 for Fhtiat. Above 60° and below 40°F the nwnber of individ-
uals with abnormal vertebrae increases. low oxygen content of water
during incubation inoreaaes the number of vertebrae.
E2;•1 rays. In counting the rays of the dorsal fin all elsnents
were inoluded. Usually. in systematioe. the mnall rays at the front of
the fin that are less than one-half the length of the longest rays are
not aounted. When oat~ating temperature effects. there is no reason
for not counting all elements. The base of all rays showed clearly in
both the rt.diographa and tho stained speoimene. but someti.'11es the longest
rays oould not be measured.
The number of dorsal rays reported for these experiments is great~r
than the number reported in the llteratW'e. Jordan and Evermarm (1896)
list 11 dor~l rays for the speoie&J Foerster and Pritchard (1935). 11
to l4J Clemans and Wilqy (1946). 10 to 14. The observed values for Ex-
periments I • II and III ranged .t'rom 13 to lB. For all three experiments
the dorsal ray oounta are recorded in Table 21 a.nd are shown graphically
in Figure 26o The ourvea are con•istent tor both the oonstant tempera-
ture and ohanging tanperature experiments with the maximum number of
raye in ·the 45° to 550F tanpt'trature range. Thie ie opposite to the ef-
teot ot temperature upon the number of Tertebrae. Tin1ng ( 1952) reported
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
109
TABU: 21
Numbers of Dorsal fur.ys for Chinook 5almon in Ex;..>a riJUent.a I, II and III
Te.mf-.1
Exp. Lot OF 13
I G 2 39.8
3 44-7
5 50.6
6 55.1
7 60.2
4 47.4*
8 4? .o
II G 3 54.6 2
5 57.8 5
4 )8. 5* 2
III Sa l 39.0 l
2 41.6
4 56.5
5 62..2 ~
6 55.5-lHI-
G l )9.0
4# 56.3
6 55.4 **
Sk 4 55.9
5
btl
61.3
54. 7**
l!; 1 38.8
.3 48.4
4 55.2
~~ bl. .3
55.2~
* At city water temperature
**At well wator temperature
•
1.4 15 16
10 91 32
34 147
27 191
9 41
3 10
18 169
32 188
ll 65 14
.56 16 1
42 37
17 17 4
1 5
12 1.40 57
8 ll
25 143 88
2 3 4
14 12
l 28 J6
14 26
11 6
8 32
14 11
3 16
1 36 1J3
1 68 86
19 159
II Water of low o.x.ygen content during incubation
17 18 .l n ~
1.5.17 133 .047
10 15.87 191 .034
33 16.02 251 .0)1
15.82 50 .05.5
14.71 13 .122
17 16.00 204 .029
14 15.92 234 .02$
14.99 92 .063
14.17 78 .062
14.43 81 .061
14.62 39 .114
14.83 6 .166
2 15.23 211 .0)8
14.43 21 .1.48
~ 15.26 258 .04(;
1.5. 22 9 .Z/8
2 15.57 28 .120
2 15.58 67 .071
1 l 15.74 42 .097
1 15.44 18 .1.45
3 15.88 43 .076
5 15.70 30 .137
2 15.95 21. .109
16 15.88 186 .039
5 1.5. 59 160 .044
46 1 16.13 225 .036
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
~
t-1 < §
~
~
§
110 16.0'
0
15.0.
14.5.
--Green
0 coostant temperature
X city water temperature
0~
40 45 50
TF}o{?ERATTJRE' ~
1s.o
1
I
I
15.5-~
I
o--
I . ---I --------15.0 t ---·· ---,.o-··
/ . .
cT
14.5 -~
o ave. of chang. temps.
x well 'Water
:14.0 1 ~ ...............
40 45 50
TEMPERATURE, 0:F
\
'
Q
b I
b II
55 60
"I-.. ~ --·. \ ..
\ ·.
\ . .
-· ·-Sacramento \ ..
--Entiat
-··Green
----Skagit
55 6o
Figure 26. .A.Terage Number of Dorsal Rays and Temperature for
lDts or Experiments I, II and III
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
111
a similar situation for the sea trout.
In oonolusion, the maximum number of dorsal rays ooourred in the
temperature range of 45° to 56°FJ this is opposite to the effect of tam-
perature upon vertebrae.
Anal rays. In counting the rays of the a.nal fin &11 elements were
included. The same &rgument for using all the elements of the dorsal
fin &lao prevails for the anal fin and~ likewiae, the number of anal
rays reported for these experiments is greater than reported in the lit-
erature.
Jordan and Evermann ( 1896) list 16 anal raya for the species. Other
authors give the following numbera S~~ultc (1931), 15 to 16s Foerster
and Pritchard (1936). 16 to 18J Clsmona and Wilby (1946), 15 to 19. The
observed values for Experiments I, II and III ranged tram 16 to 21 (for
lots other than E 8). The counts of the anr.\.1 r&\Ys are tabula.ted in Table
22 and shown graphically in Figure 27. Maximum V1Llues are in the range
of 45° to 55°F with lower values on either side of this range. For the
sea trout Tining (1952) alao found that the number of anal rays was
greatest at intennediate temperatures.
In oonolusion, as with the dorsal rays the maximum number of' anal
rays oocurred in the temperature range of 46° to 550r.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
ill
TABlE 22
Numbers of Anal Rays for Chinook Salmon of ~eriment:s I, II and III
Exp. Lot
81IlfJ. OF 16 17
I G 2 39.8
3 44.7
5 50.6
6 55.1
7 60.2 l
4 47.4*
8 47 .o
II G 3 54.6 2
5 57.8 1 38
4 58.5* 5
III Sa l 39 oO 1 ll
4 )6.5 9
5 62.2 17
6 55.5** 2l
G 1 39 .o
4 56.3
6 55.4-K-* 1
Sk 4 55.9
~~~ 61.3
54. 7**
E 1 38.8 1
3 48.4
4 55.2 1
5 61.3 15
61 55.2**
* At eit] water temperature
**At well water temtJerature
18 19 20 21
15 62 40 2
45 124 20
1 51 164 21
16 27 1
6 4
2 85 118 6
1 63 117 8
18 24
43 5
33 4
15 6
123 93 ll 1
ll 1
167 101 3
7 6
5 15 9
23 49 10
11 46 8
8 12
9 54 11 1
15 20 2
3 13 7 1
20 149 82
89 86 25 6
9 19 78 185
# Water of low oxygen content dur.ing incubation
-22 23 X n
19.24 119
19.87 189
19.86 237
19.66 44
18.27 ll
19.61 211
19.70 189
18.50 44
17.60 87
17.98 42
17.79 33
18.46 237
17.45 29
18.29 292
18.46 13
1':1.14 C1
18.82 83
18.95 65
18. &..; 20
1'/.05 75
18.61 38
19.25 24
19.24 252
18.63 221
69 5 20.82 365
s_
X
.063
.042
.036
.079
.195
.039
.040
.089
.066
.072
.136
.043
.106
.OJ6
.144
.129
. 071
.067 .
.ll2
.066
.102
.150
.038
.059
.048
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
ll3
2o.o .....
\
\
\
\ l9o0-
\
\ \
0
\ I
IB.o-·, 'T X.
---·-Green \
b II
0 constant temperatures
x ei ty water teq>eratures
17 .0.~·--~-----L-------·-·-----L--------------L-----~---
40 45 50 55 6o
20.0~
19.0 ;_ ------______.., o.. ...
_____ .-~TI
------. )( ....... ----. ----. ' ' ....
---. -""""'0 ---o-----~
----.J • •
~--· --o--··-··
. --· ,.. ' --·--· "'
11.0 ~ o a.ve. of chang. temps.
x 1iJI9ll vater
)._,. ,_ -•·· L,., . -_J._ ·-• , ..
40
-. l
-----Green
----Skagit
--Entiat
--·-Sacramento
l ...... __ ,--I-__
55 6o
Figure '2:(. Average Jlumber of Anal Ray'o~~ md Temperatures far
Lots or Ezper:1JDents r~ n and m
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
V li , 3U.YMARY
The obaervation:s from three exr.-erimen t.s U_tJLn the effects of
tem,t>erature on young chinook aa.l:nun are as follvws:
Rate of development
(1) The te;nJ.oerc.tuJ'tj coefficLents u, Q10 an·J x are not constant
for U1 e relationshi}' of te:m;.er.1 ture to the rumber of days t.o hatching.
The values for t!"E co ~L ic i.eats are ccnsider ,lJly greater at low than
LL t1igh t..em!JCr&tures wit~ a ~ritico.l te.nperatur-e i.!lvut 47°F,
\~) For lots reared <.t .;vn.stant tei:peratH't:S in 'l'"l~ ranc:;e from
J9.f30 t.o 57.8°F, tn"" t-em}erature surrunation rule, tne Lelehr&dex: equa-
tion and the losistic curve fit eyually well :..o thtl relo.tionahip of
For the te~erature summution rule, y{x-a) • k, where l • the
nwuber of deys to the time when 5L-per cent cf the e~gs are hatched.
at an incubation tem~ero. tu.re of ~ , ! is t.he t hreshcld te.Q~ra ture
and k .iB tile temt-~ratcrre sumcna.tion constant, new :net..bods for estimat-
i.nJs! and~ and a confiderx;e interv!:l.l for.! are given. Tht: value for
.Js is shown to be e . .rual to the reci}!rocal of t~ re;;ressicnar the Sfleed
of development on temperature; one estimate of!!. is XlYl-X2Y2 ; a
I Y1 -Y2
second est.imat.e of~ is i-k(:-:J; the confidence intervcJ. for the
.J
se~;ond estimate cf a is alS(., t.;iven. For the four rc.ces cf chinvok
sal.Ioon the values for ~ ran5e frv:n 31.1° to )2. 7°'i; for k, from 815
to lG20.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
The .2 value of Belehradek's equatioo ie 0.97 for the data corrected
for a threshold temperature of 33. SOF and 1.12 for the uncorrected data,
that is the threshold temperature is aeSll..ID!td to be 32°F or cfJC.
Followin~ is the equation of the logistic curve that best fits
the temperature-development relationship for lots reared at con~~t
temperatureas
- 1 + .2.46-0. 242x
y -.0396
whfl'l the incubation temperature, x, is in degrees centigrade.
(3) T~ chinook Hlmon egga from the Sacramsnto River develop 8 per
cent faster than those fran the Entiat River. The rate of develo}.lment
of the eggs from the Skagit and tile Green Rivers is intermediate.
(4) Using data from all lots rega.rdles.! of race, mortality rate,
temperature pattern, or year the equation that best fits t ru, relation-
ship of the number of days to hatching, y, and the incubation tempera-
ture, x, in degrees centigrade ia the logist-ic curve of the form
1 + e2o306-.2022x
y • .04404
for which the stands.rd error of estimate is 3 .14.
( 5) Water of low OJcy6en cootent during incubation increa.ues the
nUillber of days to hatching about 18 per cent o.t average water t~era-
turea of 55°F.
( 6) The snortel!t hatching }.leriod occurs in lots reared in the
temperature range 400 to 5f!JF for which the 5-95 percentile range is
less than five days.
( 7) Short hatching periods are associated with high survival•.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
116
(1) For the lots reared at 34°F or 65°F and higher none of the
eggs survived to the hatching stage.
( 2) vne hundred per cent mortality occurs dur:ing the yolk-sao
stage in lots reared at ti:J 0 and 62! °F.
(J) ht constant temperatures of 55° and 57~°F th~ lots hatch
successfully but during tha yolk-sac stage, mrtblity increases to
;u p .. cent or greater.
(4J The .1.ortality rate :1s low at all stages of development for
lots reared at t e:n...,eratures between 40° and 55°F.
Abnormal fry
In the temperature range 40° to 55°f the number of abnonnal fry
averagea 4. 6 J.!ai" cent par lot and at 60°r' and higher there is a nine-
fold or greater increase.
Growth
( l) At hatchine; tho fry reared at 40°f are lar<Jer than those
rea.~d at higher te.mpero.turea.
(2) The growth rate ! or lots reared at conetant temperatu:re is
greatest at 55°F and decreases in relation to the distance from the
optimum !or lots at other temperatures.
(J) For lots reared at city water temperatures, the fish are
smaller at th.; 20th weel( of the ext-er~nt than the fish reared at a
constant temperature o! 5.5oF, rut are of the same size by the 46th
week. Most ra,tJid growth occurs when the te.IIlP' rature is near 6c:P'I.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
lJ7
Weristio characters
(1) For lots reared at o~atant tem~r&tures the average number
of vertebrae is fewer in the temJ.>erature rh~e from 45° to 55°1' than
at either h~her or later temperaturee.
~2) For the ~acramento, .;;>teagit, Green and Mtiat races the number
of vertebrae average 66, o8, 69 and 72 and r~e from 63 to 78.
( 3) For lots reared at temJ.>•ratureB above 6\Pr· and below J.t:Pr·
the nwnber of individual a wit.h abnormal vertebrae increase.
( 4) ·,·t'ater of low oxyc;en content during the incubation period
increasea the average nu.ml>er of vertebrae J:-er lot as much as 2. 4.
( 5) The average nu.rrber of both dorsal and anal rays u greater
for the lot a reared in the temp~rature ron6e 45° to 55°F th8.n for lots
reared at eithdr higher or lo~r tan_t.~era.tur~. This is the opJ;Osite
of the effect of le.:ntJerature u1-on the number of vert.flbrae,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
LITERATURE CITED
.~DRE't'iA..~THA, H. G. and L. C. Birch
1954 The distribution and abundance of animals
Chicago, University of Chicago rress. 782 ~lJ·
A.f'STEIN, C.
1909 Dilj beetimmUfl6 des Alters pelagisch 1ebender Fis cheier,
~ittoilunger des Deutschen Seefischerei-Vereins, 25(2):364-373.
Translated by .W. Crull.
,.. '
n.E.LEHRADEK, J.
1929 Sur la signification de5 c~fficient.s de temy.;rature.
rrotoplasma, 7(2;:232-255. Translated by U. ~~rriman.
::lELEHRADEK, J.
1935 Tamp~rature and livi% ;natter.
1'rotclJ1a.sma .i.ionogrt.q nien 8 (Berlin). 277 d·.
bl!;LDING, D. L., M. J. l-ender and J. 11., Rodd
19j2 'l'h~ early 5rowth of salmun _l...larr in Ganadi<ill hbtcheries.
Trans. Ml. Fishuries .:Joe., 62:2.11-2~3.
BG;m;..~, .1\. and ·~;. H. Bay li.f f
195) hadiography of a.!Dbil fisll: .s for meristic studies.
BRETT, J.
1952
CoJ-eia, 1953(3) :150-15L
R.
Tem_f;erc.ture t c1er on:::e in young 1-'acif ic sal.n~n, genus
Uncorhynchua.
J. Fiah:Iies Research Board. ~an., <t(o):265-J2J.
bi\L, WN lN v, T • L •
1'752 The influence cf temf·.=rc.tl..U'e on the rate of develo~nt of
insect~, with S!JdCial refuran~e t0 the et,e;;s of vry1lulus
COfiltil(ldUS "rib.lker.
~ustralian J. 3ci. Res., d5:'15-lll.
BUhNE.R, C. J.
11.51 Charact~istics oi spawnint rests of CoL:mbia "'ivur salmon.
LI. S. Fish Wildlife .Service, F.ishcry Bull., 52( 61) :97-110.
BUF' •. ho~·,·s, R.
1956 MS.
CLE.\iliNS,
1946
'ii. ;.... and G. V. Wilby
Fiohes of the 1-'acific cOb.st of Canada.
Fisheriee Research Board Can., Bull. 68, 368 Pfi.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
CLOTHIER,
1950
ll9
c. R.
A key to same southern California rishes based on vertebral
ohare.ctar a.
Calir. Fish Game, Fish. Sullo 79. 83 pp.
COPE, O. B.
1949 Water temperature records fran Calirornia' s Central Valley,
1939·1948.
u. s. Fiah Wildlife Ssrvioe, Speoial Scientific Reporta Fish-
eries No. 2, 77 PP•
COPE, o. B.
1952 water temperatures of California's Central Valley, 1949-51.
COSTE, J.
1858
CREASER,
1930
U. s. Fiah Wildlife Servioe, Speoial Soientific Reportz Fish-
eries No. 75, 48 PP•
J,.
Instructions pratiques aur la pisiou1ture.
2nd ed. Paris. 144 PP•
Cited from Hayes, 1949.
c. w.
Relative importance or hydrogen-ion conoerrtration, temperature,
dissolved oxygen, and carbon~diaxide tension, on habitat seleo-
tion by brook-trout.
Ecology, ll(2)a246-262.
DANNEVI G, A.
1960 The inrlu$noa of the environment on number of vertebrae in
plaioe.
Fiskeridirektoratets Skrifter (rteports on Norwegian Fishery
and Marine Investigation), Sar1e Havunderspkelser, 9(9)a6 pp.
DANNEVIG, H.
1895 The 1nfluanoe of temperature on the development of the eggs of
fishes.
Soot1and Fishery Board (Glasgow), Ann. Rep. for 1894, Scienti-
fic Invest., l3(8)al47-l52.
DAVIDSON I J.
1944 On the relationship betTI8en temperature and rate of develo~ent
of inseota at constant tanp8l"aturaa.
DAVY, J.
1856
J. Animal Eool., 13(l)t26•33.
On the vitality of the ova of the Salmonidae or different ages.
Proo. Royal Soo. (London), 8e27~3.
DONALDSON, J • R •
1950 A problem on the relation of oold temperatures to the develop-
ment of sockeye salmon eggs.
MS.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
12C1
DONALDSON, J. R.
1955 Experimental studies on the aurv1 val of the early stages of
chinook salmon a1'ter carrying exposures to upper lethal tem-
peratures.
A thesis submitted for the degree of Master of Soienoe, Seattle,
University of Washington. 116 PP•
DONALDSON, L. R. and F. Foe tel'
1940 Experimental study of the effect of various water temperatures
on the growth, food utili~tion, and mortality r~tes of f1ngsr-
ling sockeye aa~on.
Trans. Am. Fisheries Soc., 70a3~9-S46.
ELLIS, C. H.
1953 Correspondence of August, 1953.
ELLlS, M. M., B. A· Westfall and M. D. Ellis
1948 Determination of water quality.
u. s. Fish Wildlife Service, Research Report 9, l22 pp.
EMBODY,
1934
G. C.
Relation of t~perature to the incubation periods of eggs of
four species of trout.
Trans. Am. Fisheries Soo., 64t28l-292.
FALLERT, s.
1952 Interview, October 13, 1952.
FO~STER,.
1936
R. E. and A· L. Pritchard
A study of the variation in certain meristic
genus Onoorhynohus in British Columbia.
Trans. Roy. Soc. Can.,Seotion V,. pp. 85-95~
characters in the
FOSTER, R. F., et al.
1949 The effect on embryos and young of rainbow trout frcm exposing
the parent fish to X-rays.
Growth, 13sll9-l42.
GABRIEL, M. L.
1944 Fa.otors affecting the nl1Ilber t.nd form of vertebrae in Fundulus
heteroo 11 tua.
J. Exp. Zool., 95t~05-145.
GILL, T. N.
1863 A note on the Labro1ds of the western ooast of North America.
GINSBURG,
1938
Proo. Aoa.d. N'at. Soi. (Philadelphia), l5a22l-222.
I.
Arithmetical definition of the species, sub-species and race
concept with a proposal for a modified nomenclature.
Zoologioa,. 231253-286.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
GRAY, J.
1928
GR:r;EN, S.
1870
" GUNT~,
1862
The growth of fish. III. The effect of tsmperature on the
development of the eggs of Salmo fa.ri o.
Brit. J. Exp. Biol., 6(2)al25-1!o.
Trout oulture.
RochetSter, New York, Preae of Curtis Morey &: Co. 90 PP•
A.
Catalogue of the fiehea in the British Yuseum.
British ~useum (Natural History), Zoolo~y Department (London),
4a5:54 PP•
HAYES, F. R.
1949 The growth, ~enera.l chemistry and temperature relations of
salm.on1d eggs.
REINCKE,
1898
HEl!T S, lo!.
1947
Quart. Rev. Biol., 24(4)•281-308.
F.
Naturgesohiohte des Herings. Tell I. Die Lokalformen und die
" ~nderungen des HerinGG in den europaisohen Meeren.
Abhandlungen des Deutachen Seefisoherei-Vereins, 2(l)al-238~
Cited fram Schmidt, 1917.
J.
Expertmental studies on adaptive evolution in Gasterosteus
aoulea tus L.
Evolution, lr89-l02.
HEU'l' S , M • J.
1949 Racial divergence in fin ray variation patterns in G~sterosteus
aouleatus.
J. Oeneto, 49(:5).183-191.
HOLLISTER, G.
1934 Clearing and dyeing fish for bone study.
Zoologioa, l2a89-10l.
HOURSTOU, W. R.
19&3 Correspondence of August 12, 1953.
HUBBS, C.
1925
L.
Racial and seasonal variation in the Paoif1c herring, Califor-
nia aardine and California anchovy.
Calife Fiah Game, Fish. Bull. a. 2~ PP•
JACKSON, R.
195~ Correspondence of July 17, 1953.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
: ..., . .,
JANISCH, E,. II
" 1925 Uber die tamperaturabhangigkeit
kurvamatzige ana1vse. " ., ,,
biologisoher Vorgange und deren
JOHANSEN,
1914
JOHNSON,
1953
Pflugers Arohiv. i\ir di~;"J gesarnte Physiologie d. Mensohen u. d.
Tiere No. 209, PP• 414-436.
A. c. and A· Krogh
The ini'luenoe ot temperature and certain other factors upon
the rate of development or the eggs of fiahea.
Publ. Ciro. Cons. Explor. Mer. 68a44 PP•
H. E. and R. F. Brice
Effects of transportation of green ege;a and ot water tanpera-
ture during incubation on the mortality of chinook salmon.
Progressiv~ Fish Cu1turist, 15(3)al04-108.
J<EDAN, D. S.
1891 Relations of ter:1perature to vertebrae among fishes.
Proo. u.s. Nat. Mus., l4al07-120.
JORDAN, D. S • and B • W. Everma.nn
1896 The fishes of North Qnd Middle Americ~~
Bull. 1!. s. Nat. Mus., 47, Part I, 1240 PP•
KA YiA.JIR I ,
l327a
M.
The optimum temperature of water for the hatching of the eggs
of trout, Oncorhynchus ma5o-u (Walbt.Uin). _
J. Imp. Fisheries Inst. (Japan), 23(2)al4-19.
KAWAJIRI, M.
l927b The influence of variation of tamperature of water on the de-
velo~ent of f'ish~e;cs.
KAWAJIRI,
1928
KROOH. A.
1914
LINDSEY,
1954
J. Imp. Fisheries Inst. (Japan), 4S(3)•66-7~.
M.
The influence of variation ~f temperature of water on the de-
velopment of fish-eggs. II.
J. Imp. Fisheries Inst. (Japan), 24 ( 1) al-4.
On the influence of the temperature
development., z. Allg. Physiol., l6al63-177.
Cit,d fran Hayes, 1949.
c. c.
on the ra. te of embryonic
Temperature-controlled meristio variation in the paradise fish
Ma.oropodus opercularia (L.).
Can. J. Zoof., 52(2)a87-98.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
MARCDLUlN, K.
1964 Is there any correlation between metabolism and number of
vertebrae (and other meristic characters) in the sea trout
( Salmo trutta trutta L.) f
Keddeleaer fra Danmarks Fiskeri, og FAvundera~gelser Ny
Serie, l(~)al-9.
MATTSON, C. R.
194t8 Spawning ground studies of Wilhm.tte River spring ohinook
salmon.
Oregon Fish. Comm. Res. Briefs, l(2)a21-32.
MoHUGH, J. L.
1954& The influence of light on the numb•r of vertebrae in the grun-
ion, Leuresthea tenuia.
Copeia, 1964(1)a2~-26.
lloHWH. J. L.
l954b Geographic variation in the Paoifio herring.
Copeia, l954(2)el39-l~l.
»ERRDUJI • D •
19~5 The effect of temperature on the development of the eggs and
larvae of the out-throat (Salmo o!Arldi olarkii Richardson).
J. Exp. Biol., 12(4)J297-S05 •
.MOFFSTT,
1949
J. w.
The first four years of king-salmon maintenance below Shasta
D~, Saer~ento RiTer, Californi&c
Calif. Fish Game, 36(2)t77-102.
MCYrTLEY, C. KoC •
19~ The eff&Ot of t-.mperature during development on the number of
aoales in the Kualoopa trout, S&lmo kamloops Jordan.
Contrib. can. Biol. and Fiaher!ea, nvs., 8(2o)a254-263.
KOTTLBT, c. KoC.
1937 Tho number of vertebrae in tro\lt (Salao).
!ORRIS.
1868
T.
J. Biol. Board Can., 3(Z)al89-l16.
AJnerioan fiah culture, embracing all the detaile o!' artificial
breeding and re&r'ing of trouta the oult\U"e of salmon. shad and
other fishes.
Philadelphia, Porter and Coates. S04 pp.
OLSOH, P. A. and R. F• Foster
1956 Temperature toleranoe of eggs and young or Columbia River chin-
ook ae.lmon.
To be published in Volume 85 of the Trans. ~. Fisheries Soo.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PEARL, R.
1920
and L. J. Reed
On the rate o! growth of the population of the United States
ainoe 1790 and ita math.matioal repre•entetion.
Pro. Nat. Aoad. Soi. Uft s • ., 6(6)a276-288.
PELNAR, J.
1953 Correepondenoe of Nov~ber 17, 1953o
PRICE, J. l'f.
1940 Time-temperature relatiex!s in the inoubation of the whitefish.,
CoregGnue olupeaformia (Mitohill)~
J. Gen. Phyaio1., 2!(4)a449-468.
I
RE.AlJllUR 5
l7S8
REIBISCH.,
1902
ROCX19EIL,
1966
R, A. F.
Obaervations du thermametre fa.ites a. Paris pendent l•a.rwee
MDCCXXV.
Memoires de Mathematique et de Physique Annee 1735, pp. 545-
576. (In Hiatoire de l•Aoademie Royale des Soienoea, Paris).
Cited tram Thompson, 1952.
J.
Ueber den Einflues der Temperatur auf die Entwiokelung von
Fisoh Eiern.
Wisaensohaftliohe Meereeuntersuohungen. Kiel (abt. Kiel),
n.a., 6(2)a213-231.
J.
Sane effects of' sea water and temperature on the embryos of'
the Paoifio almon, Onoorhynohua gorbusoha (Walba.um) and Onoor-
hynohus keta ('ifalbaum).
I theils--.ubmitted in partial fulfillment for -!;he der;ree
Doctor of Philosophy, Seattle., Univ.rsity of Washingtono
of
416 pp.
ROUNSE.FELL, G, A. and E. H. Dahle;ren
19~2 Fluctuations in the tupply of h~ing, Clupea pallasii, in
Prince W1111Ul Sour:xl, Alaaka.
Bull. U. s. Bur. Fish •• 47(9)t263-291.
RUCK~, P.. R.
19~7 The effect of temperature on the growth or the embryo of Oncor-
~chua nerk:a (Wa.lbtlum).
heals submitted for the degree of Master of Scienoe, Seattle,
University of washington. 64 pp.
SCIDIIDT,
1917
J.
I. Zoe.roee vi Vi F.• L. and local
Lee Canptes-Ren usdes Travaux du
279-396.
raoes of the same,
Laboratoire Carlsberg, 13(3)z
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
SCl:DtlD'l',
l919a
scmo: DT '
l9l9b
SCHMID'l',
1920
125
J.
Racial studiea in fishes.
with Lebiatea retioula.tua
Jo Genet •• e(s}.1~1-15S.
J.
Racial atudiea in fishes.
(S&~o trutta L.)~
J. Genet •• 9(1)a61-67.
J.
II. EXperimental investigations
(Peters) Regan.
III· Diallele crossings with trout
Racial inveatigationao IV. Experimental investigations with
Zoa.roea vivi~aroua L•
J. Genet., 1 {!)el79-191.
SCBMII11', J.
1921 Racial investigations. VII. Annual tlucrtuatione ot racial
oharaoters in Zoaroes vh'i.parous.
Las Camptes-Rendus des tra.vaux du Labora.toire Carlsberg, 14(15)•
24 ppe
SCHMIDT, J.
1930 Rao1a.l investigations. x. The Atla.ntio Cod (Gadus eallaria.s
L.) and looal ra.oes of the a&me.
Les Camptes-Rendus des Travaux du Laborato1re Carlsberg, 18(6)z
71 PP•
SCHULTZ, L. P.
1931 Key to the fishes of W&ahington and Oregon.
Seattle, University Bookstore. 63 pp.
SEATTU
1963
&.i'ER WORKS DE?!' •
SET.at!OOR,
1963
SHELFORD,
1927
SMITH, S.
1960
Annual report of the W&ter
for the year 1953. 64 PP•
A. H. and J c R. Donaldson
Departm.nt ot the City of Seattle
occurrence of high mortality among chinook salmon fi'y after
the recharge ot a charcoal filter.
Progresaive Fiah Cultur1at, l6(S),l29-1SO.
V. E.
An experimental investigation of the relations of the oodling
moth to weather and olimateo
Illinois Natural History Survey Bull., l6•Sll-440 (Art. 5).
H.
Upper Sacramento River sport fishery.
u. s. Fish Wildlife service, Special Scientific Reports Fish-
eries No. 34~ 44 PP•
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
126
A. V.
:Kxperimenta em meristic and other oharaoters in fishes. I. ().1
the influenoe.of temperature on same meristic oharaoters in
sea-trout and the function period of these oharaoters.
Medd. Iamm. Havandua. Serle Fiskeri, 11(3)a66 PP•
!iWING~ A. V.
1946 st~ge of determination of vertebrae in teleostean fishes.
0
'!.UliiD,
1960
Nature (London), 157(~992),694-596.
A. V.
Influence of th$ environnent of number of vertebrae in tele-
oatean fishes.
Nature (London), l65(,184)a28.
Tflo:OO, A. Vo
1952 Experimental study of meristic character• in fishes.
Biol. Rev. Cambridge Phil$ Soc., 27(2)al69-193.
TESTER, A. L.
1938 Variation in the mean vertebral count of herring (Clupe& pal-
1aaii) with l'l'&ter temporature.
J. Conae Int. Explor. Mer., l3(l)a7l-75.
THOMPSON, D1 A. W.
1952 on growth and torm.
2nd ad. (1942), ~eprinted 1952.
Cambridge, England, University Presso Vol. 1, 466 pp.
TOWNSEND, L. D.
1944 Variation in the DUmber of pyloric caeca and other numerical
oha.raotere in chinook salJilon and in trout.
Cope!~, 1S~4(1)a52-54.
UVAROV, B. P.
19~1 Inaeots and ~1Lmate.
Trana. R. Ent. soc. London, 79(1)a1-247.
VLADYKOV, V • D •
1964 Te.xonood.o ohara.oters of the ea3tern North America chars (~
v.linua and CriatoYOm8r).
J. Flih. Research Board Can., ll(6),9~-932o
VOUTE. A. D.
1936 z. Angew. Entom •• 22al-25.
Cit~d from Davidson, 1944.
ULLICH, C.
1901 A method o~ reoordin~ egg dsvelopment. for use of fish oultur-
ists.
u. S. Bureau of Fia.lleriea, llf!.shington, D. Co Report of the
Commi1aioner of Fisheries for 1900, Doo. 452, PP• 187-194.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
l27
WEISEL, G. Fe
1955 Variations in tM number o£ !'in raya of two oyprinid fishes
oorrelated with natura.l water temperatureao
soology, 36(l)al-6.
'WELANDER, A. D., et &1.
1948 The etreotor Roentgen rays on the embryos and larvae of' the
ohinoolc salmon.
Growth, l2(3)t20S-242.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
VITA
Allyn HenrJ seymour was born ir1 Seattle. Waal:ington. on August l, 1913.
HilS parents are Alphonse Shel"W'ood S~r and Velma. Smith Se~:monr.
He attended ~ High School in Seattle and entered the University
of Waahingt::m in 1932 . The requirements for the degree of Bachelor of Science
in F~sheries were c~leted in December 1937.
Since 1937 he hs£1 worked with the International Fisheries Comnrl.ssion,
the International Pacific Salmon Fisheries Canrm:ission and the Wa.sh:i.n[:ton State
Depa.riment of Fisher:t.es. Em;ployment with the Applied Fisheries Laboratory,
University of Washin.0ton, befan in 1947 and has continued to the present.