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University of Southern California Dissertations and Theses
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Physiology and biochemistry of food limitation in marine invertebrate larvae
(USC Thesis Other)
Physiology and biochemistry of food limitation in marine invertebrate larvae
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Content
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iv
Acknowledgements
How to begin? There have been so many people who have contributed to making this
dissertation possible and my life in Los Angeles bearable.
In Hollywood fashion, "I'd like to thank the members of the Academy..." my thesis
dissertation committee Prof. Donal Manahan, Prof. Dennis Hedgecock, Prof. Steven Finkel and
Prof. Robert Maxson. My patient dissertation committee members provided oversight through this
enormous undertaking, and made sure resources and financial support were available for my
education and research. Prof. Hedgecock was especially helpful for all the resources from our
laboratory collaborations, from genotyping manpower to qPCR; Chapters 1 and 4 would've been
impossible without the advice and resources of the Hedgecock lab group.
I must give thanks to the late, great Bill Trusten, who welcomed me to the campus, lent his
ear and unfailingly helped me navigate the bureaucracy of being a student. Linda Bazilian has not
only filled his shoes, but taken on an even more supportive role for which I am eternally grateful.
Don Bingham has made sure that the administrative paperwork of the department and its resources
were readily available to me.
I can't thank Wrigley Marine Science Center enough. They were my summer home for 6
years and they took me in as one of them. The staff were so awesome and supportive to me and
more importantly, patient. Lauren Garske and Lauren Czarnecki were fantastic as friends and lab
managers; their support was invaluable for the success of Chapters 1 and 2. I have to especially
thank Maureen Oudin for logistic support, Steve McElroy for not killing me every time I made a
mess of the third floor, and Randy Phelps for bringing a beautiful companion tarantula into my
v
life. I made such great friends working out there, especially Lisa Collins, Laura "Dancing Queen"
Jordan for the rays, Annaliese Hettinger, Matt Iacchei for the lobsters, and Patrick Robinson for
comaraderie. And finally Dr. Kathy Ann Miller, who opened her home and her library to me, and
who was an allimportant link to my hometown, provided me many hours of wisdom and advice.
On campus, I've gotten invaluable support from scientists in the department. Prof.
Michelle Arbeitman and Justin Dalton provided assistance and LSCM time. Prof. Michael Quick
provided advice about transporter biochemistry, reagents for my experiments, and invaluable
insight into life in academia, and into life in general. Alicia Thompson did all the sectioning for
the TEM work, and provided training and encouragement. Sangeeta Bardhan Cook was my protein
biochemistry cheerleaderwithout her logistical support and expert advice, I could not have
completed Chapter 3. I hope I can support you as much as you did me. And thanks to Jason
Duncan and Prof. John Tower for training and access to the original confocal where most of the
microscopy was done.
My former Manahan lab mates have been like a second family to me. Dr. Douglas Pace
and Dr. Jason Curole were my academic big brothers, prodding me always to become a better
scientist, like them. Dr. Eli Meyer provided samples and for guidance for Chapters 3 and 4, and as
a core member of the field team, pitched in with culturing when I needed an extra set of hands. Dr.
Allison Green, in addition to helping with training me and providing culturing assistance, was a
trailblazer in the lab and paved the way for me. She had the determination and gumption to make
it to the other side of the barrier, like a larva crossing a steep halocline.
I couldn't have had a better gang of Los Angeles friends, inside and outside the program.
Julie Rose opened her home to me and helped make me an Eastside gal; her energy and
vi
professionalism were a great inspiration and example. Dave Ehrlich and Sheri Ozeki were my
gateway friends in LA, who took me in immediately and introduced me to so many of their
friends, their community, and their favorite things about the city. Adriane and Max, and the Aqua
House posseKeren, Mahira and Dianethank you for the great times. I'll miss you so much.
George Rose, you are fabulous. I especially have to thank my closest friends who stuck by me,
before, during and through to the end. Prof. Scott Kroken and Dr. Eleanor Kincaid were role
models and friends, without whom I wouldn't have made it to grad school. Matt Cornell stuck by
me, lending an ear when I most needed it, and gave me a cultural life to counterbalance the science.
Dr. Steven Flusty provided love, magick and spiritual encouragement. Thank you for lighting the
path in the darkness. And lastly Dr. Paul Bayman, who's love and patience through the years
provided the stability I'd never had before, and who always reminded me how much fun science
could be.
Funding for my tuition and research was generously provided by an Oakley Fellowship
from the College of Letters and Science at USC, the Rose Hills Summer Fellowships from the
USC Wrigley Institute for Environmental Studies, a Final Summer Dissertation Completion
Fellowship from the Graduate School at USC, and most importantly, the National Science
Foundation.
vii
Table of Contents
Epigraph ................................................................................................................................................................ ii
Dedication ........................................................................................................................................................... iii
Acknowledgements ........................................................................................................................................... iv
List of Tables ...................................................................................................................................................... ix
List of Figures ...................................................................................................................................................... x
Abstract .............................................................................................................................................................. xiii
Introduction ........................................................................................................................................................ 1
Introduction References ........................................................................................................................... 19
Chapter 1: Genotype variation in survival and poststarvation growth recovery of bivalve
larvae Crassostrea gigas (Thunberg, 1793) .......................................................................................... 32
Abstract ...................................................................................................................................................... 32
Introduction ................................................................................................................................................ 34
Methods ....................................................................................................................................................... 38
Results .......................................................................................................................................................... 44
Discussion ................................................................................................................................................... 60
Chapter 1 References ............................................................................................................................... 72
Chapter 2: Starvation physiology, and growth recovery to metamorphosis of echinoderm
larvae Lytechinus pictus (Verrill 1867) ................................................................................................ 82
Abstract ...................................................................................................................................................... 82
Introduction ................................................................................................................................................ 84
Methods ....................................................................................................................................................... 86
Results .......................................................................................................................................................... 89
Discussion ................................................................................................................................................ 105
Chapter 2 References ............................................................................................................................ 115
Chapter 3: Localization and expression of ectodermal amino acid transporters in the early
development of the purple sea urchin Strongylocentrotus purpuratus (Stimpson 1857) ..... 122
Abstract ................................................................................................................................................... 122
Introduction ............................................................................................................................................. 124
Methods .................................................................................................................................................... 126
Results ....................................................................................................................................................... 135
Discussion ................................................................................................................................................ 159
Chapter 3 References ............................................................................................................................ 171
viii
Chapter 4: Sequencing and ontogenic expression of putative amino acid transporters from
larvae of the Pacific Oyster, Crassostrea gigas (Thunberg, 1793) ............................................... 177
Abstract ................................................................................................................................................... 177
Introduction ............................................................................................................................................. 179
Methods .................................................................................................................................................... 180
Results ....................................................................................................................................................... 185
Discussion ................................................................................................................................................ 199
Chapter 4 References ............................................................................................................................ 206
References ....................................................................................................................................................... 210
Appendix A: Supplement to Chapter 1 .................................................................................................... 234
Appendix B: Supplement to Chapter 4 .................................................................................................... 239
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% Survivorship from day 2
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b17E#T!&A>#`>#$>#&-5#e#cG')*1D)-(#MMM>#B'')-51G#B#T3''4)D)-(&4#b17E#T!;A>#a#&-5#M#B'')-51G#B#
T3''4)D)-(&4#b17E#T!9AE#])&-#(.(&4#41'15#9.-()-(#1-#"6!#5&/#.45#0)417)*2#8&2#!EY#l#VE+#-7#&9*.22#
:&D141)2>#&99.3-(1-7#:.*#O"!R#.:#(%)#.*7&-19#D&22#*&-7)#+Y6"@REA##N)'419&()#934(3*)2#.:#:&D14/#
B#%&5#9.-212()-(#41'15#9.-()-(#&(#5&/#"#!E"#-7#4&*0&
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A#&-5#&(#5&/#+V#+EQ6+E?#-7#4&*0
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217-1:19&-(#9%&-7)2#1-#(.(&4#41'15#9.-()-(#.-)68&/#BjP^B>#'KVE!AE#=*1&9/474/9)*.42#&99.3-()5#
:.*#&.3(#"@R#.:#(%)#5&/#"#41'15#9.-()-(#.-#&0)*&7)>#;3(#8&2#+VR#;/#5&/#!>#&-5#8&2#3-5)()9(&4)#
!
W+#
:*.D#5&/#W#.-8&*5#1-#&44#:&D141)2E#$%&-7)2#1-#2()*.4#9.-()-(#53*1-7#2(&*0&(1.-#8)*)#3-5)()9(&4)#
53)#(.#(%)#2D&44#&D.3-(2#.:#2()*.4#VE+6VE"W#-7#4&*0&
6+
AX#41'15#9.-()-(#&:()*#5&/#W#9.-212()5#
'*1D&*14/#.:#'%.2'%.41'15#O@WRAE##
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;)(8))-#:&D141)2#81(%#*)4&()5#'&*)-(2#8)*)#.:#1-()*)2(E#T3*010.*2%1'#.:#*)4&()5#4&*0&)#%&4:621;41-7#
:&D141)2#81(%#(%)#15)-(19&4#D&()*-&4#'&*)-(#:*.D#41-)#oY+o>#cG')*1D)-(#MMMA#%&5#0)*/#51::)*)-(#
23*010&4#(1D)2#1-#.-)#)G')*1D)-(#9.D'&*1-7#(8.#4&*0&4#:&D141)2#b17E#+E"AE#=%12#'&1*#.:#:&D141)2#
51::)*)5#1-#23*010&4#(1D)#;/#+EJ#:.45#&(#(%)#WVR#23*010.*2%1'#4)0)4>#&-5#;/#"E+#:.45#&(#(%)#+R#
23*010.*2%1'#4)0)4E#=%)#4.7#(*&-2:.*D#23*010&4#*&()2#8)*)#%17%4/#217-1:19&-(4/#51::)*)-(#BjP^B#.:#
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&-5#&-#&0)*&7)#(1D)#(.#+R#23*010.*2%1'>#8%14)#:&D14/#`#%&5#D)51&-#++E+#5&/2#&D.-7#+J#
:&D141)2A#(1D)#(.#WVR#23*010.*2%1'#++EW#5&/2A#&-5#&-#)G9)'(1.-&44/#4.-7#(1D)#(.#+R#23*010.*2%1'>#
1-519&(1-7#*)4&(10)#5)9*)&2)#1-#D.*(&41(/#81(%#&7)E#P0)*#WVVV#4&*0&)#OVEWR#23*010&4A#8)*)#2(144#
&410)#&(#5&/#!Y#1-#:&D14/#`#&**.8AE#=%)2)#(8.#:&D141)2#51::)*)5#217-1:19&-(4/#(6()2(>#'pVEVWA#1-#
(.(&4#'*.()1-#9.-()-(#=&4)#+E+A#&-5#-.(#217-1:19&-(4/#1-#(.(&4#41'15#9.-()-(#QE"#-7#4&*0&
6+
#02#!EJ#
-7#4&*0&
6+
>#B'')-51G#B#T3''4)D)-(&4#b17E#T!;A#&(#5&/#"X#%.8)0)*#(%)*)#8&2#-.#217-1:19&-(#
51::)*)-9)#;)(8))-#(%)#(8.#:&D141)2#1-#'*.()1-#9.-()-(#&(#5&/#+Q>#5)2'1()#(%)#+EY6:.45#51::)*)-9)#1-#
23*010.*2%1'E#N)2'1*&(1.-#*&()2#1-#;.(%#:&D141)2#=&4)#+E+A#8)*)#-.(#217-1:19&-(4/#51::)*)-(#.0)*#(%)#
9.3*2)#.:#(%)#')*1.5#.:#:..5#5)'*10&(1.-#.-)68&/#BjP^B>#'KVEVWAE#
#
!
W"#
b173*)#+E"#
Age (days)
0 5 10 15 20 25 30 35 40
% Survivorship from day 2
0
20
40
60
80
100
B-35 x 81
C-51 x 81
#
b173*)#+E"E#I&*7)#23*010.*2%1'#51::)*)-9)2#;)(8))-#3-:)5#%&4:621;41-7#4&*0&)#:*.D#:&D141)2#`#&-5#$#
cG')*1D)-(#MMMAE#I&*0&)#5)0)4.')5#:*.D#7&D)()2#.:#.-)#41-)#Y+#:)D&4)>#8)*)#:)*(141L)5#81(%#2')*D#
:*.D#(8.#51::)*)-(#D&4)2#41-)#W+67*)/#&-5#!W69321JN'&"%"-2($%&'(I,'*024"'"&&/')23#621;41-7#4&*0&4#
:&D141)2#`#!W#G#Y+A#&-5#$#W+#G#Y+AE#I&*7)#51::)*)-9)2#8)*)#.;2)*0)5#1-#23*010.*2%1'#5)2'1()#(%)#
2%&*)5#D&()*-&4#'&*)-(E#B**.8#1-519&()2#VEWR#23*010.*2%1'#1-#:&D14/#`#!W#G#Y+AE##
#
#
#
#
!
W!#
])(&.419#;357)(##
B#'.()-(1&4#)G'4&-&(1.-#:.*#)G()-5)5#23*010.*2%1'#1-#(%)#&2)-9)#.:#:))51-7#9.345#;)#)1(%)*#
&*7)#*)2)*0)#)-)*7/#23''4/#:*.D#D&()*-&44/623''41)5#;1.D&22>#.*#D)(&.419#5)'*)221.-#(.#
D1-1D1L)#(%)#9.2(#.:#'%/21.4.719&4#D&1-()-&-9)E#M-#.*5)*#(.#1-0)2(17&()#(%)#&22.91&(1.-#;)(8))-#
23*010&4#&-5#;1.9%)D12(*/#8)#4..C)5#:.*#9.**)4&(1.-2#;)(8))-#'*.()1-#&-5#23*010.*2%1'>#&2#(%)#
*)4&(10)4/#2D&44#O"VRA#'*.'.*(1.-#.:#41'15#1-#(%)2)#4&*0&)#12#4)22#41C)4/#(.#9.-(*1;3()#(.#23*010&4#
0&*1&(1.-#&(#(%12#5)0)4.'D)-(&4#2(&7)E#=%)#*)4&(1.-2%1'#;)(8))-#&0)*&7)5#(.(&4#'*.()1-#9.-()-(#
.0)*#(%)#53*&(1.-#.:#2(&*0&(1.-A#&-5#(1D)#(.#WVR#23*010.*2%1'#8&2#-.(#217-1:19&-(#*
"
U#VEVY>#
'KVE!AE#S1(%#1-9*)&21-7#D.*(&41(/>#(%)#9.**)4&(1.-#;)(8))-#&0)*&7)5#'*.()1-#9.-()-(#&-5#(1D)#(.#
+R#23*010.*2%1'#8&2#8)&C4/#217-1:19&-(#&-5#5)')-5)-(#.-#(8.#)G9)'(1.-&4#:&D141)2#81(%#0)*/#%17%#
23*010.*2%1'#&-5#&.0)#&0)*&7)#'*.()1-#9.-()-(#*
"
UVE!@>#'pVEVWA#b17E#+E!&AE#=%)#*&-7)#.:#1-1(1&4#
'*.()1-#9.-()-(2#&(#5&/#"#=&4)#+E+A#9.0)*)5#.-4/#*&-7)#.:#"EW#:.45#51::)*)-9)>#2417%(4/#4)22#(%&-#
(%)#*&-7)#.:#+R#23*010.*2%1'E#=%)*)#8&2#-.#217-1:19&-(#9.**)4&(1.-#;)(8))-#')*9)-(#'*.()1-#4.2(#
&-5#23*010&4#(1D)#*
"
UVE+!>#'KVE!#:.*#(1D)#(.#+RAE#B0)*&7)5#*)2'1*&(1.-#*&()2#53*1-7#:..5#
5)'*10&(1.-#515#-.(#9.**)4&()#81(%#(1D)#(.#+R#23*010.*2%1'#b17E#+E!;g##*
"
UVEV?>#'KVE!A#.*#WVR#
23*010.*2%1'#*
"
UVEVQ>#'KVEQAE#
,*.8(%#*)9.0)*/#
# b.*#3-:)5#4&*0&)#&4)#(.#23*010)#'&2(#(%).*)(19&4#HjN>#(%)#&141(/#(.#*)9.0)*#&-5#7*.8#1-#
(%)#'*)2)-9)#.:#:..5#8.345#&42.#;)#)22)-(1&4#:.*#:3*(%)*#23*010&4E#].*&-#&-5#]&-&%&-#"VVQA#
2%.8)5#(%&(#'.2(62(&*0&(1.-#*)9.0)*/#.993*2#1-#(%12#2')91)2E#,*.8(%#*)9.0)*/#:*.D#2(&*0&(1.-#8&2#
9.-212()-(4/#.;2)*0)5#&2#1-9*)&21-7#2%)44#4)-7(%A#23;2)_3)-(#(.#(%)#1-(*.539(1.-#.:#:..5#1-#(8.#W+#
G#!W#:&D141)2#2%.8-#cG')*1D)-(2#M#&-5#M^>#b17E#+EQAE#N)9.0)*/#.:#4&*0&)#81(%#1-9*)&21-7#')*1.52#
!
WQ#
b173*)#+E!E#`1.9%)D12(*/#&-5#'%/21.4.7/#9.D'&*)5#81(%#(1D)#(.#+R#23*010.*2%1'E#BA#=.(&4#'*.()1-#
9.-()-(#&0)*&7)5#.0)*#934(3*)#53*&(1.-#02E#(1D)#(.#+R#23*010.*2%1'E#c**.*#;&*2#*)'*)2)-(#TEcE]E#:.*#
D34(1'4)#2&D'4)#'.1-(2E#*
"
UVE!@>#'pVEVWE#`A#B0)*&7)5#*)2'1*&(1.-#*&()2#.0)*#934(3*)#53*&(1.-#02E#
(1D)#(.#+R#23*010.*2%1'E#c**.*#;&*2#*)'*)2)-(#TEcE]#.0)*#D34(1'4)#2&D'4)#'.1-(2#.*#TEcE]E#.:#
24.')#:.*#934(3*)2#(%&(#8)*)#D)&23*)5#.-4/#.-9)E#*
"
UVEV?>#jTE#=%)#*&-7)2#.:#*)2'1*&(1.-#*&()2#&-5#
23*010&4#(1D)2#&*)#;.(%#&''*.G1D&()4/#Q6:.45E#
!
WW#
b173*)#+E!#
Total protein (ng larva
-1
)
4 6 8 10 12 14 16
Time to 1% survivorship (days)
5
10
15
20
25
30
35
40
#
Respiration rate (pmol O
2
larva
-1
hr
-1
)
0 5 10 15 20 25
Time to 1% survivorship (days)
5
10
15
20
25
30
35
40
B
A
r
2
=0.39
r
2
=0.07
!
WJ#
#
b173*)#+EQ#
Age (days)
0 10 20 30 40 50
Shell length ( m)
80
100
120
140
160
180
200
220
51 x 35-IA
51 x 35-IB
51 x 35-IC
51 x 35-I control
51 x 35-IV
51 x 35-IV control
#
#
b173*)#+EQE#H.2(62(&*0&(1.-#7*.8(%#*)9.0)*/#1-#(8.#:&D141)2#.:#4&*0&)#cG')*1D)-(2#M#&-5#M^AE#
])&-#21L)#.:#4&*0&)#kDA#lTc#53*1-7#2(&*0&(1.-#&-5#53*1-7#:))51-7#*)9.0)*/#&2#:..5#8&2#
1-(*.539)5#&(#51::)*)-(#(1D)2E#$.-(*.4#(*)&(D)-(2#;4&9C#&-5#7*)/#51&D.-52A#8)*)#:)5#&(#5&/#"#&-5#
D)&23*)5#5&14/E#T%)44#4)-7(%#D)&23*)D)-(2#-UWV#&(#)&9%#'.1-(A#:*.D#:)5#;*)&C.3(#934(3*)2#.:#(%)#
W+#G#!W6M#4&*0&4#:&D14/#cG')*1D)-(#Mg##B>#`>#&-5#$#*)'419&()#934(3*)2#&*)#*)'*)2)-()5#;/#:144)5#
2/D;.42>#*)2')9(10)4/>#&2#1-#b17E#+E+;A>#&-5#(%)#W+#G#!W6M^#4&*0&4#:&D14/#cG')*1D)-(#M^g##7*)/#
91*94)2>#2))#&42.#b17E#+EWAE##
!
W?#
.:#:..5#5)'*10&(1.-#8&2#D)&23*)5#:.*#+6"#8))C#')*1.52>#&-5#*)9.0)*1-7#4&*0&)#7*)8#(.#21L)2#
9.D'&*&4)#(.#9.-(*.4#4&*0&)#:)5#&(#5&/#"A#&:()*#&-#1-1(1&4#4&7#')*1.5>#3'#(.#QV#5&/2#.:#1-1(1&4#:..5#
5)'*10&(1.-E#=%)#(8.#W+#G#!W#:&D141)2#()2()5#:.*#2(&*0&(1.-#*)9.0)*/#&44#23*010)5#3-:)5#:.*#.0)*#"J#
5&/2#:&D14/#W+#G#!W6M#:*.D#b17E#+E+;X#&-5#:&D14/#W+#G#!W6M^#1-#b17E#+EW&A>#&-5#7*.8(%#.:#
*)9.0)*1-7#4&*0&)#1-#&44#934(3*)2#8&2#217-1:19&-(E#
,*.8(%#%)()*.212#&-5#23*010.*2%1'#
# =.#9.D'&*)#7*.8(%#'.()-(1&4#&-5#23*010&4#'.()-(1&4#81(%1-#-74)#:&D14/>#9.%.*(2#8)*)#
2)'&*&()5#1-(.#:)5#&-5#2(&*0)5#(*)&(D)-(2#cG')*1D)-(#M^AE#BD.-7#(%)#:.3*#4&*0&4#:&D141)2#:*.D#
(%)#:&9(.*1&4#9*.22#.:#41-)2#!W#&-5#W+>#(8.#:&D141)2#%&5#0)*/#51::)*)-(1&()5#23*010&4#*&()2#8%14)#(%)#
*)D&1-1-7#(8.#:&D141)2#%&5#1-()*D)51&()#23*010.*2%1'#b17E#+EW&AX#&*0&4#:&D14/#5)*10)5#:*.D#
*I$356(6+"/'9-,,50(,1J'$0'0),I%'#,-'1,4+2-$0,%'2%5')20'0$4$32-'0.-A$A,-0)$+'(,'#24$36'BW#G#W+E#
=%12#W+#G#!W#:&D14/#%&5#(%)#%17%)2(#23*010.*2%1'#.:#&-/#:&D14/#&D.-7#&44#)G')*1D)-(2#.')-#
(*1&-74)2>#2))#&42.#:&D14/#W+#G#!W6M^#1-#b17E#+EQA#&-5#8&2#"6:.45#D.*)#2(&*0&(1.-#*)212(&-(#(%&-#1(2#
*)91'*.9&4#:&D14/#!W#G#W+X#(%12#:&D14/#&42.#%&5#7*)&()*#4.-7)01(/#(%&-#;.(%#1(2#%&4:621;41-7#:&D141)2#
!W#G#!W#&-5#W+#G#W+#81(%#)1(%)*#2%&*)5#)77#.*#2')*DAE##
=%)#4.7#(*&-2:.*D#.:#23*010&4#*&()#2%.8)5#(%&(#(%)#cG')*1D)-(#M^#:&D141)2#8)*)#
217-1:19&-(4/#51::)*)-(#BjP^B#.:#24.')2>#'pVEVV+A#&-5#(%&(#(%)#24.')2#8)*)#217-1:19&-(4/#
51::)*)-(#;/#:&D14/#;&2)5#.-#(%)#,I]#BjP^B#.-#9.-(*&2(2>#'pVEVV+AE#=%)#W+G!W#:&D14/#8&2#
217-1:19&-(4/#7*)&()*#(%&-#(%)#D15'&*)-(#23*010&4#*&()#fW!KV#:.*#%)()*.212>#'pVEVV+A>#;3(#(%)#
!WGW+#8&2#-.(#217-1:19&-(4/#51::)*)-(#:*.D#(%)#D15'&*)-(#*&()>#f!WU#V#:.*#&551(101(/>#'KVEQAE#M-#
(%)#'.2(6%.9#()2(2>#.-4/#W+G!W#8&2#217-1:19&-(4/#%17%)*#(%&-#(%)#o%17%#'&*)-(o#:&D14/#!WG!W##
!
WY#
b173*)#+EWE#j.-6&551(10)#(*&1(2#1-#Q#:&D141)2#.:#4&*0&)#cG')*1D)-(#M^A#:.*#5&/#J#9.-(*.4#7*.8(%>#
2(&*0&(1.-#*)212(&-9)#&-5#*)9.0)*/#7*.8(%E#BA#e1::)*)-(1&4#23*010.*2%1'#.:#:.3*#4&*0&4#:&D141)2#
7)-)*&()5#;/#'&*)-(#./2()*2#:*.D#(8.#;*..52(.9C#:&D141)2#W+#&-5#!W7'8'*I$35(6+"/'1-,00'$0'0),I%'
:.*#9.D'&*12.-E#=%)#W+#G#!W##24$36'K230,'*C+#G#!W6PQ/'$%'R$&7'+EQA#%&2#217-1:19&-(4/#51::)*)-(#
2(&*0&(1.-#*)212(&-9)#BjP^B#.-#24.')2>#'pVEVV+A#9.D'&*)5#(.#1(2#*)4&()5#:&D141)2#!W#G#!W>#W+#
G#W+>#!W#G#W+AE#1-2)(A#T%)44#4)-7(%#kDA#&(#5&/#J#&-5#7*.8(%#*&()#kD#5&/
6+
A#.:#21;41-7#4&*0&)#:.*#
(%)#2&D)#:.3*#:&D141)2#(%&(#8)*)#:)5#.-#5&/#"#:&D14/#W+#G#!W>#024"'20'*C+#G#!W6PQ'1,%(-,3/'$%'
b17E#+EQA#1-2()&5#.:#;)1-7#2(&*0)5E#B44#7*.8(%#*&()2#&*)#217-1:19&-(4/#51::)*)-(#:*.D#)&9%#.(%)*#(8.6
8&/#BjP^B#'pVEVV+X#`.-:)**.-1#'ppVEVVY!#:.*#&44#'&1*812)#9.D'&*12.-2AE#`A#N)9.0)*/#
7*.8(%#.:#:.3*#:&D141)2#:)5#&(#5&/#+YE#,*.8(%#*&()2#1-2)(A#&*)#217-1:19&-(4/#51::)*)-(#;)(8))-#
:&D141)2#BjP^B#.-#24.')2>#'pVEVV+AE#c**.*#;&*2#:.*#TEcE]E#.:#24.')#41)#81(%1-#(%)#
*)'*)2)-(&(1.-2#.:#(%)#5&(&E#
!
W@#
b173*)#+EW#
Age (days)
0 10 20 30 40 50
Percent survivorship from Day 2
0
20
40
60
80
100
35 x 35
51 x 35
35 x 51
51 x 51
wild-type
Family
Growth rate ( m day
-1
)
0
5
10
15
Shell length ( m)
100
120
140
160
35x35
51x35
35x51
51x51
A
Age (days)
0 18 21 24 27 30 33
!"#$$%$#&'("%)*+,
80
100
120
140
160
180
200
220
35 x 35
51 x 35
35 x 51
51 x 51
Family
-./0("%.1(#%)*+%213
-1
)
3
6
9
12
B
51x51
51x35
35x51
35x35
#
!
JV#
%"+EV>#BjP^B#.-#'.2(6%.9#9.-(*&2(2>#'pVEVWA>#&-5#!WG!W#8&2#217-1:19&-(4/#7*)&()*#(%&-#!WGW+#
BjP^B#.-#'.2(6%.9#9.-(*&2(2>#'pVEVWAE#
=%)#9.-(*.4#:)5A#4&*0&)#8)*)#D)&23*)5#5&14/#:*.D#5&/#"#3-(14#5&/#J>#&-5#(%)#W+G!W#:&D14/#
%&5#(%)#4&*7)2(#21L)#&(#5&/#J#]&--6S%1(-)/#F#()2(>#'pVEVV+A>#&-5#(%)#%17%)2(#7*.8(%#*&()#(8.6
8&/#BjP^B>#'pVEVV+A#.0)*#:.3*#5&/2#.:#:))51-7#b17E#+EW-2)(AE#,*.8(%#*)9.0)*/#3'.-#
:))51-7#&(#5&/#+Y#8&2#.;2)*0)5#1-#&44#:&D141)2#:*.D#cG')*1D)-(#M^#b17E#+EW;AE#B44#:.3*#7*.8(%#
*&()2#8)*)#217-1:19&-(4/#51::)*)-(#BjP^B#.-#24.')2>#'pVEVV+>#b17E#+EW;#1-2)(A#&-5#8)*)#(*)&()5#
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Age at 50% survivorship (days)
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A
B
Age (days)
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Lipid content as % of total egg lipid
0
20
40
60
80
100
Family
1 3 6
Total
Phospholipids
Triacylglycerols
Sterols
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7'59'()/=#3'8B)(#56B)5/#()1()/)39#90)#3'8B)(#.2#46;/#.2#2))4%3.(#90)#7.(()/1.34%3/'(-%-.(/0%1+#
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#
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Age (days)
0 10 20 30 40
Percent survivorship from day 3
0.5
5
50
1
10
100
Family 2
Family 5
Family 6A
Family 6B
#
#
Age at first feeding (days)
3 7 11 15 20 24
Percent survival with12 days of feeding
0
20
40
60
80
100
7 18
Family 1
Family 3
Family 6 B
Family 6 A
#
A
B
9
6
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#
Age (days)
0 10 20 30 40
Total lipid (ng larva
-1
)
0
50
100
150
200
Family 6A-starved
Family 6B-starved
Family 6B-Fed@3 (control)
Family 6A-Fed@11
Family 6B-Fed@20-1
Family 6B-Fed@20-2
#
#
#
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#
#
#
Age (days)
0 5 10 15 20 25 30 35
Protein mass (ng larva
-1
)
0
100
200
300
400
500
600
Family 3-starved
Family 3-Fed@7
Family 1-Fed@7
Family 1-Fed@18
Family 1-starved
Families 1 & 3
# Family 6
Age (days)
0 10 20 30 40
Protein mass (ng larva
-1
)
0
100
200
300
400
Family 6A-starved
Family 6B-starved
Family 6B-Fed@3 (control)
Family 6A-Fed@7
Family 6A-Fed@11
Family 6B-Fed@15
Family 6B-Fed@20-1
Family 6B-Fed@20-2
Family 6
A
B
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Age (days)
0 10 20 30 40
Respiration (pmol O
2
larva
-1
hr
-1
)
0
100
200
300
400
Family 1 Fed@7
Family 3-starved
Family 3 Fed@7
Family 1 Fed@18
Family 1-starved
0 10 20
0
5
10
15
20
$$
$
$$ $
$
Family 1 Fed@18
at day 44
A
B C
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Age (days)
0 5 10 15 20 25
% Biomass energy available
-25
0
25
50
75
100
125
Energy equivalents ( J)
0
500
1000
1500
2000
2500
theoretical energy remaining
actual biomass energy
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A
Sp-AT3
Sp-AT2
Sp-AT1
250
160
105
75
50
35
30
Total Protein per lane
0.1 1 10 100
Relative area units
0
2000
4000
6000
8000
B
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Age (days)
0 3 6 9 12 15 18
Protein (ng larva
-1
)
0
20
40
60
80
100
120
A
embryo
A
B
C
D
E
F
Age (days)
4 6 8 10 12 14 16
Transport (pmol Ala larva
-1
hr
-1
)
0
5
10
15
20
25
30
35
A
B
C
D
E
F
B
Total protein (ng larva
-1
)
20 40 60 80 100 120
Transport rate (pmol Ala larva
-1
hr
-1
)
5
10
15
20
25
30
35
Prot
12 15 18 21 24
Trans
4
5
6
7
8
C
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Age (days)
0.0 0.5 1.0 1.5 2.0 2.5 3.0
Relative Units per lane
0
1000
2000
3000
4000
A
Sp-AT1
Sp-AT2
Sp-AT3
$
$
Relative Units
per larva
B
Age (days)
5 10 11 15
0
50
100
150
200
Starved
Fed
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Relative Units
per larva
0
7
14
21
28
35
C
Age (days)
5 10 11 15
Starved
Fed
$
Age (days)
Relative Units
per larva
0
20
40
60
80
D
5 10 11 15
Starved
Fed
$
$
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1 2 3 4 5 6 7 8 9 10 11 12
1 2 3 4 5 6 7 8 9 10 11 12
100 200 300 400 500 600 700
Hydropathy Score
-3
-2
-1
0
1
2
3
4
Amino Acid Position
0 100 200 300 400 500 600 700
Hydropathy Score
-3
-2
-1
0
1
2
3
4
A
B
Amino acid position
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#
#
A B
Larval family
35x35 51x35 35x51 51x51
Fold expression of Cg-AAT1 0.1
1
10
a
a,b
c
b
Larval family
35x35 51x35 35x51 51x51
Shell length ( m)
80
85
90
95
100
105
110
115
a
a
b
c
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Age
0 10 20 30 40 50
Mortality
0.0
0.2
0.4
0.6
0.8
1.0
5 x 3
5 x 3 regression
5 x 5
5 x 5 regression
3 x 5
3 x 5 regression
3 x 3
3 x 3 regression
A
Age
0 10 20 30 40 50
Ln [survivorship x 100]
-1
0
1
2
3
4
5
3 x 3
5 x 5
3 x 5
5 x 3
20% survivorship
Truncated linear regression
B
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Culture or treatment
FSW(0.22) FSW(1.0) 5x5 48 h 5x3 0 h 5x3 12h 5x3 36h 5x3 48 h
Bacterial abundance (x 10
6
ml
-1
)
0
2
4
6
8
10
before or control
after or treatment
*
*
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Age (days)
0 2 4 6 8 10 12 14 16
Total lipid (ng larva
-1
)
0
1
2
3
4
5
F2-A
F2-B
F2-C
A
Age (days)
0 2 4 6 8 10 12 14 16
Total lipid (ng larva
-1
)
0
1
2
3
4
5
3x8
5x8
8x5
B
Age (days)
0 2 4 6 8 10 12 14 16
Total lipid (ng larva
-1
)
0
1
2
3
4
5
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$
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Age (days)
1 2 3 5 10 14 18 109
Expression Ratio
0.01
0.1
1
10
100
1000
Age (days)
1 2 3 5 10 14 18 109
Expression Ratio
0.01
0.1
1
10
100
B
A
35x35-2
51x35-2
51x51-2
35x51-2
midparent
35x35-1
51x35-1
51x51-1
35x51-1
midparent
$
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2 3 5 10 14
0
100
200
2 3 5 10 14 18
0
100
200
Shell length (µm) Shell length (µm)
35x35-1
35x51-1
51x35-1
51x51-1
35x35-2
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Age (days)
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Expression Ratio
0
1
2
3
4
Age (days)
1 2 3 5 10 14 18 109
Expression Ratio
0
1
2
3
4
5
C
D
3x3-2
5x3-2
5x5-2
3x5-2
3x3-1
5x3-1
5x5-1
3x5-1
$
Abstract (if available)
Abstract
For high fecundity organisms, the number of offspring that survive to adulthood are very few. The mechanisms of individual-based survival remain of great interest in biology. In this dissertation, larval survival was examined by taking an integrative approach, from organismal physiology to molecular biology.
Linked assets
University of Southern California Dissertations and Theses
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Asset Metadata
Creator
Yu, Pauline Chia-Wen
(author)
Core Title
Physiology and biochemistry of food limitation in marine invertebrate larvae
School
College of Letters, Arts and Sciences
Degree
Doctor of Philosophy
Degree Program
Biology
Degree Conferral Date
2009-08
Publication Date
07/29/2009
Defense Date
05/07/2009
Publisher
University of Southern California
(original),
University of Southern California. Libraries
(digital)
Tag
amino acid transporter,energy budget,larvae,OAI-PMH Harvest,pluteus,starvation recovery,starvation survival,veliger
Language
English
Contributor
Electronically uploaded by the author
(provenance)
Advisor
Manahan, Donal T. (
committee chair
), Finkel, Steven E. (
committee member
), Hedgecock, Dennis (
committee member
), Maxson, Robert E. (
committee member
)
Creator Email
pcy@usc.edu,splene@gmail.com
Permanent Link (DOI)
https://doi.org/10.25549/usctheses-m2402
Unique identifier
UC1442230
Identifier
etd-Yu-2966 (filename),usctheses-m40 (legacy collection record id),usctheses-c127-567328 (legacy record id),usctheses-m2402 (legacy record id)
Legacy Identifier
etd-Yu-2966.pdf
Dmrecord
567328
Document Type
Dissertation
Rights
Yu, Pauline Chia-Wen
Type
texts
Source
University of Southern California
(contributing entity),
University of Southern California Dissertations and Theses
(collection)
Repository Name
Libraries, University of Southern California
Repository Location
Los Angeles, California
Repository Email
cisadmin@lib.usc.edu
Tags
amino acid transporter
energy budget
larvae
pluteus
starvation recovery
starvation survival
veliger