An experimental examination of temperature interactions in the ‘match-mismatch’ hypothesis for Pacific cod

larvae

Benjamin J. Laurel, Thomas P. Hurst, and Lorenzo Ciannelli1Fisheries Behavioral Ecology Program, NOAA RACE Division-Alaska

Fisheries Science Center, Newport, OR 973652College of Oceanic and Atmospheric Sciences, Oregon State University,

Corvallis OR, 97331-5503

Pacific cod

-Rank 2nd in catch and product value in the groundfish fishery in Alaska (much smaller relative component of fishery in Canada)

-Pacific cod has been examined in a multi-species framework e.g., regime shifts, climate change and North Pacific trophodynamics

-Few studies have explicitly examined the ecology and life history of Pacific cod possibly because of assumed similarities between Pacific cod and their better-studied congener, Atlantic cod (Gadusmorhua).

Sea ice coverage in the Bering sea has changed over the past 30 years

0

500

1000

1500

2000

2500

3000

78 80 82 84 86 88 90 92 94 96 9820

0020

0220

04

Biom

ass

(100

0 mt)

050100150200250300350400450500

Catch

YC Age3 Millions

Age 3+Bio

Climate affects the ecosystem through sea ice

February March April May June

Slide courtesy of Gary ShawHunt et al. 2002

The Oscillation Control Hypothesis contradicts predictions based on the

‘Match/mismatch’ hypothesis

Month

Feb Mar Apr May Jun Jul Aug

Abu

ndan

ce (n

o./m

2) o

f lar

vae

in p

ositi

ve to

ws

0

20

40

60

80

100

120

140 Peak Pacific cod larval production (1972-1996)Yr 1997 primary productivity (sea-ice present)Yr 2000 primary productivity (sea-ice absent)

OCH or match-mismatch? Complex interactions

Prey field(timing, magnitude)Temperature

field

Post-feeding stages(growth and survival)

Pre-feeding stages(hatch duration,

endogenousresources)

Modeling growth and survival of Pacific cod eggs/larvae in response to climate-related changes in sea ice conditions in

the Bering Sea

• Phase I: Vital rates of eggs and pre-feeding larvae in response to temperature

• Phase II: Vital rates of post-feeding larvae and juveniles in response to interactions in temperature and food availability

• Phase III: Development of spatially explicit models of early life survival of Pacific cod in the Bering Sea using laboratory validated data

Temperature controlat the NOAA-FBEP laboratory at the HMSC in Newport, OR

Flow-through, temperature controlledseawater system

Experiment #1: Pre-feeding larvae

• 5 temperatures (0, 2, 4, 6 and 8ºC)– Larvae: 3 replicate containers for each hatching

stage: ~25%, ~50% and 75% hatch (n=45 tanks)

• Measured daily mortality and morphometricchanges (yolk area, MH, SL and ED) every 3 days to ~50% mortality

• Lipid/fatty acid characteristics measured at early, mid and late hatch for each temperature treatment

Hatching is temperature-mediated

0 2 4 6 84

8

12

16

20

24

0 2 4 6 8 1010

20

30

40

50

60

Temperature (ºC)

Dur

atio

n (d

ays)

Hatch Duration Timing to Hatch

0° C

020406080

100

2° C

020406080

100

Early hatchMid- hatchLate hatch

4° C

X Data

% S

urvi

val

020406080

100

6° C

X Data

020406080

100

8° C

DPH

0 10 20 30 40

020406080

100

Early hatchers survive longeracross all temperature treatments

4° C

Days post hatch0 10 20 30 40

% S

urvi

val

0

20

40

60

80

100

Early hatchMid hatchLate hatch

Temperature

0 2 4 6 8 10

DPH

to 5

0% m

orta

lity

(LO

G+1

)

0.9

1.0

1.1

1.2

1.3

1.4

1.5

1.6Early HatchLate Hatch

Size and growth SL

Days into hatch cycle

0 2 4 6 8 10

Size

(SL

mm

)

4.2

4.4

4.6

4.8

5.0

5.2

5.4

5.60 degrees2 degrees 4 degrees6 degrees8 degrees

Temperature

0 2 4 6 8 10

Max

siz

e (S

L m

m)

4.9

5.0

5.1

5.2

5.3

5.4

5.5

5.6

8°C

6°C

4°C

2°C

Day 3 - P. cod larvae

0°C

0° C

8° C

62 DPF

The consequences of cold and warm environments

LF-LF

Experiment #2: Post-feeding larvae

6 wks3ºC

8ºC

HF-HFLF-LFHF-LFLF-HF

NF

6 wksHF-HF

HF-LFLF-HF

NF

Temperature mediated match-mismatch experiment

Match-mismatches in prey at two temperatures

• Temperature explains the most variance in growth

• Fish at cold temperatures survived longer in the absence of prey

• The relative timing of match-mismatches in prey therefore differed between temperature treatments

Week0 1 2 3 4 5 6 7

SL

(mm

)

5

6

7

8

9

10

11

12HFHF HFLF LFHF LFLF

Week0 1 2 3 4 5 6 7

SL

(mm

)

5

6

7

8

9

10

11

12HFHFHFLFLFHF LFLF

3ºC

8ºC

8ºC

Week0 1 2 3 4 5 6 7

SL

(mm

)

5

6

7

8

9HFHF HFLF LFHF LFLF

3ºC

Week0 1 2 3 4 5 6 7

SL

(mm

)

5

6

7

8

9

10

11

12HFHFHFLFLFHF LFLF

8ºC

Growth consequences of early and late mismatches of prey

Conclusions• Time of hatch and hatch cycle duration are negatively correlated

with temperature—early hatchers survive longer in the absence of prey but may have a limited ability to feed on certain sizes and types of prey relative to late hatchers.

• Optimal conversion efficiency using endogenous reserves occurs at colder temperatures (≤4ºC). Larvae grow larger and survive longer in the complete absence of prey at colder temperatures

• (3ºC) Mismatches in food availability at cold temperatures are relatively inconsequential at low temperatures except in the complete absence of food. However, early and late mismatches in food have measurable effects on growth

• (8ºC) Mismatches in food availability are very important to growth, especially when mismatches occur after 3 wks post-hatch.

• Temperature ultimately drives growth when sufficient prey for survival are present

Future and ongoing work

• Further integration into Bering sea models (Phase III)

• The behavioral consequences of early and late hatching in a predator and feeding environment

AcknowledgementsFundingNorth Pacific Research Board (NPRB - Grant#: R0605, NOAA Fisheries – Alaska Fisheries Science Center RACE Division

Co-PIs: Al Stoner-NOAA/AFSC and Mike Behrenfeld-OSU

PersonnelField assistance: Alisa Abookire and Brian Knoth

Boat charters: Tim Tripp and Jan Axel

Laboratory experiments: Scott Haines, Michele Ottmar and PaulIseri

Match-mismatch results wk 6 at 8ºC

• Optimal feeding conditions contribute to highest growth

• Late mismatches in food are more consequential to growth than early mismatches.

Match-mismatch results wk 6 at 3ºC

• Optimal feeding conditions matter little in terms of growth

• Late mismatches still appear to be consequential to growth compared to early mismatches.

Survival% survival at wk 6

Feeding conditions

HFHF LFLF HFLF LFHF

% S

urvi

val

0

10

20

30

40

50

60

708 degrees 4 degrees

Match-mismatch results wk 6 at 3ºC

Preliminary model runs

1999(coolest)

2005(warmest)

Experiment #2: Post-feeding larvae

• 2 temperatures (3 and 8ºC)• 6 wk experiment with 5 feeding treatments

per temperature: HF-HF, LF-LF, HF-LF, LF-HF and no food. Three replicate incubators per food-temperature treatment (n=30 tanks)

• Morphometric measurements using image analysis taken every wk (SL, MH, ED, Yolk area)

• % survival measured at the end of the experiment in the food treatments. Daily mortality recorded for the no food treatments

Questions?

Changes in lipid classes at 2ºCTAG

DPF

5 10 15 20 25 30 35

µg li

pid/

egg

or la

rvae

0.0

0.2

0.4

0.6

0.8

1.0

1.2

PL

DPF

5 10 15 20 25 30 35

µg li

pid/

egg

or la

rvae

0

2

4

6

8

10

12

AMPL

DPF

5 10 15 20 25 30 35

µg li

pid/

egg

or la

rvae

0.0

0.1

0.2

0.3

0.4

0.5

0.6

Sterols

DPF

5 10 15 20 25 30 35

µg li

pid/

egg

or la

rvae

0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

Changes in total lipid composition of early and late hatching eggs as a function of

temperature

Days post fertilization

5 10 15 20 25 30 35

µg li

pid

3

4

5

6

7

8

9

10

112°C4°C6°C8°C

8°C

6°C

2°C

Day 3 - P. cod larvae

0°C

4°C

Egg collection March 2006

Coulson et al. 2006

Similar but perhaps not as similar as was previously thought……

Pacific cod

Atlantic cod

0º C

DPH

0 5 10 15 20 25 30

Siz

e (S

L m

m)

4.6

4.8

5.0

5.2

5.4

5.6

DPH0 5 10 15 20 25S

ize

(SL

mm

)4.6

4.8

5.0

5.2

5.4

5.6

EarlyMidLate

DPH0 2 4 6 8 101214161820

Siz

e (S

L m

m)

4.24.44.64.85.05.25.45.6

DPH0 2 4 6 8 10 12 14

Siz

e (S

L m

m)

4.4

4.6

4.8

5.0

5.2

5.4

DPH0 2 4 6 8 10 12 14

Siz

e (S

L m

m)

4.54.64.74.84.95.05.15.25.3

2º C 4º C

6º C 8º C

Length

Eye diameter0º C

DPH

0 5 10 15 20 25 30

Eye

dia

met

er (m

m)

0.24

0.26

0.28

0.30

0.32

0.34

0.36

DPH0 5 10 15 20 25

Eye

dia

met

er (m

m)

0.270.280.290.300.31

0.320.33

0.34

EarlyMidLate

DPH0 2 4 6 8 101214161820

Eye

dia

met

er (m

m)

0.24

0.26

0.28

0.30

0.32

0.34

DPH0 2 4 6 8 10 12 14

Eye

dia

met

er (m

m)

0.28

0.29

0.30

0.31

0.32

0.33

0.34

DPH0 2 4 6 8 10 12 14

Eye

dia

met

er (m

m)

0.27

0.28

0.29

0.30

0.31

0.32

0.33

2º C 4º C

6º C 8º C

Condition Index0º C

DPH

0 5 10 15 20 25 30

Con

ditio

n (M

H/S

L m

m)

0.0460.0480.0500.0520.0540.0560.0580.0600.062

DPH0 5 10 15 20 25

Con

ditio

n (M

H/S

L m

m)

0.0460.0480.0500.0520.0540.0560.0580.060

EarlyMidLate

DPH0 2 4 6 8 101214161820

Con

ditio

n (M

H/S

L m

m)

0.0460.0480.0500.0520.0540.0560.0580.0600.062

DPH0 2 4 6 8 10 12 14

Con

ditio

n (M

H/S

L m

m)

0.0460.0480.0500.0520.0540.0560.0580.0600.0620.064

DPH0 2 4 6 8 10 12 14

Con

ditio

n (M

H/S

L m

m)

0.0480.0500.0520.0540.0560.0580.0600.0620.0640.066

2º C 4º C

6º C 8º C

Yolk depletion0º C

DPH

0 5 10 15 20 25 30

Yol

k ar

ea (m

m2 )

0.0

0.1

0.2

0.3

DPH0 5 10 15 20 25

Yol

k ar

ea (m

m2 )

0.000.050.100.150.200.250.300.35

EarlyMidLate

DPH0 2 4 6 8 101214161820

Yol

k ar

ea (m

m2 )

0.00

0.05

0.10

0.15

0.20

0.25

0.30

DPH0 2 4 6 8 10 12 14

Yol

k ar

ea (m

m2 )

0.00

0.05

0.10

0.15

0.20

0.25

0.30

DPH0 2 4 6 8 10 12 14

Yol

k ar

ea (m

m2 )

0.000.050.100.150.200.250.300.35

2º C 4º C

6º C 8º C

Pacific cod

-Rank 2nd in catch and product value in the groundfish fishery in Alaska (much smaller relative component of fishery in Canada)

-Pacific cod has been examined in a multi-species framework e.g., regime shifts, climate change and North Pacific trophodynamics

-Few studies have explicitly examined the ecology and life history of Pacific cod possibly because of assumed similarities between Pacific cod and their better-studied congener, Atlantic cod (Gadusmorhua).

Lipid catabolism in Pacific cod eggs and pre-feeding larvae

Lipid catabolism in Atlantic cod eggs and pre-feeding larvae

Days post fertilization

1 4 7 9 11 14

µg li

pid

/ egg

or l

arva

0

2

4

6

8

10

12Total lipidPhospholipidsTriacylglycerol Hatch

Lipogenesis in Pacific cod eggs/larvae across all temperature treatments

0 Degrees

Days post fertilization0 10 20 30 40 50

µg li

pid

/ egg

or l

arva

0

1

2

3

4

5

6

7

2 Degrees

Days post fertilization0 5 10 15 20 25 30 35

µg li

pid

/ egg

or l

arva

0

2

4

6

8

10

12

4 Degrees

Days post fertilization0 5 10 15 20 25 30

µg li

pid

/ egg

or l

arva

0

1

2

3

4

5

6

7

8

6 Degrees

Days post fertilization0 2 4 6 8 10 12 14 16 18 20

µg li

pid

/ egg

or l

arva

1

2

3

4

5

6

7

8 Degrees

Days post fertilization0 2 4 6 8 10 12 14 16

µg li

pid

/ egg

or l

arva

0

2

4

6

8

10

Changes in total lipid composition of early and late hatching eggs as a function of

temperature

Days post fertilization

5 10 15 20 25 30 35

µg li

pid

3

4

5

6

7

8

9

10

112°C4°C6°C8°C

Yolk area

Hatch day0 2 4 6 8 10

Yol

k ar

ea (m

m2 )

0.00

0.05

0.10

0.15

0.20

0.25

0.30

0.350 degrees2 degrees 4 degrees6 degrees8 degrees

2°C

4°C

8°C

6°C

0°C