Outline
Introduction
Temperature and Sex
Egg Water Exchange
Eggs and Dirt: The Vertebrate Seed Bank
Ralph A. Ackerman
Department of Ecology, Evolutionary and Organismal Biology
March 5, 2009
1 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
1
Introduction
2
Temperature and Sex
Soil Temperature: A Problem
A Dose Model
Temperature Cycling
Dose Response
Climate
Soil Temperature
Soil Temperature and Dose
3
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
2 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
TSD
Introduction
3 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Green Turtle Eggs
4 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Green Turtle Eggs
5 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Green Turtle Eggs
6 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Green Turtle Eggs
7 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Green Turtle Eggs
8 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Turtle Embryo Hatching
The Egg (Hatching)
Egg Shell
9 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Turtle Embryo Hatching
The Egg (Hatching)
Egg Shell
Egg Shell Membrane(s)
10 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Turtle Embryo Hatching
The Egg (Hatching)
Egg Shell
Egg Shell Membrane(s)
Egg Tooth on Hatchling
11 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Green Turtle Eggs
12 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Research Agenda
Topics
Exchange of water, heat and respiratory gases between
embryos, eggs and the incubator
13 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Research Agenda
Topics
Exchange of water, heat and respiratory gases between
embryos, eggs and the incubator
Effect of exchange on embryonic development
14 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Research Agenda
Topics
Exchange of water, heat and respiratory gases between
embryos, eggs and the incubator
Effect of exchange on embryonic development
How the incubator works (i.e. what is the climate in the seed
bank)
15 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
TSD
Temperature and Sex
16 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Constant Temperature Experimental Design
Sex Ratio (% male or female)
0.1 1
10
40 70
95 99.5
100
70
Elapsed Time (days)
60
50
40
30
20
10
0
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
o
Incubation Temperature ( C)
17 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Hatchling Sex and Incubation Temperature
1.0
Chrysemys picta
Sex Ratio (% female)
0.8
0.6
0.4
0.2
0.0
18
20
22
24
26
28
30
32
34
36
38
o
Temperature ( C)
18 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Shifting Temperature Experimental Design
Sex Ratio (% male or female)
0.1 1
10
40 70
95 99.5
100
70
Shift Once
Elapsed Time (days)
60
Shift Twice
50
40
30
Sensitive
Period
20
10
0
20 21
22 23 24
25 26 27
28 29 30
31 32 33
34 35
o
Incubation Temperature ( C)
19 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
TSD and Reptiles
Reptiles and TSD
All sea turtles
All crocodilians
Some but not all
freshwater turtles
Some lizards
No Snakes
20 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Fermi Model and TSD
1.0
Chrysemys picta
Sex Ratio (% female)
0.8
0.6
0.4
Tpiv= 28.10
0.2
a = -0.63
SE = 0.15
SE=0.15
0.0
18
20
22
24
26
28
30
32
34
36
38
o
Temperature ( C)
21 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Fermi Model and TSD
22 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Summary of Tpiv Data
23 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
TSD and Reptiles: Our Model Organisms
24 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
TSD
Soil Temperature: A Problem
25 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Soil Temperature at Ames
25
o
Temperature ( C)
Ames, 2003
20
15
Mean Soil Temperature (C)
5 cm 19.06
10 cm 18.90
20 cm 18.04
50 cm 15.97
100 cm 14.09
10
185
190
195
200
205
210
Julian Day
26 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Chelydra Nest Temperature
Nest Temperature
40
o
Temperature ( C)
35
30
25
20
Top
Bottom
15
10
200
202
204
206
208
210
212
214
216
218
220
Julian Day
40
o
Temperature ( C)
35
30
25
20
15
10
170
180
190
200
210
220
230
240
250
260
Julian Day
27 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Chelydra Nest and Soil Temperature
28 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
TSD
A Dose Model
29 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Double Shift (Daily) Experimental Design
o
Dose about 12 hours
32
Incubator
o
Temperature ( C)
30
28
Egg Box
26
24
6 hours
22
20
18
0
400
800
1200
1600
2000
2400
Time of Day (hrs)
30 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Response: Take 1
Mn ’96
Ia ’96
Sex Ratio (% Female)
1.0
0.8
0.6
0.4
0.2
Bars are 95% CI calculated for
binomially distributed data
0.0
0
o
Constant 25 C
5
10
15
20
25
30
o
Daily Dose ( hrs)
31 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Response: Take 2
Mn ’96
Ia ’96
Mn ’97
Ia ’97
Sex Ratio (% Female)
1.0
0.8
0.6
0.4
0.2
Bars are 95% CI calculated for
binomially distributed data
0.0
0
o
Constant 25 C
5
10
15
20
25
30
o
Daily Dose ( hrs)
32 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Response Curve: Take 3
33 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Thermal Dose and Incubation Period
34 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
TSD
Soil Temperature Climate
35 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Snapping Turtle Range and Climate Stations
36 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Painted Turtle Range and Climate Stations
37 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Weather Stations
38 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Mean 70 Day Incubation Period Temperatures
39 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Response Curve: Take 4
40 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Weather Stations
41 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Summer Dose Data-Hours Above 29
42 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Response Surface-July
43 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Response: Take 5
44 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
A Dose Model
Climate
Dose Seasonal Variation at Ames
45 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
TSD
Egg (Seed) Water Exchange
46 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Sea Turtle Nesting
47 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Sea Turtle Nesting
48 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Sea Turtle Nesting
49 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Sea Turtle Nesting
50 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Standard Egg Incubation-Laboratory
Box + medium + gas phase
51 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Standard Egg Incubation-Laboratory
Box + medium + gas phase
52 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Standard Egg Incubation-Laboratory
Box + medium + gas phase
53 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Incubation in Nature
Egg Clutch Completely Buried in Soil
54 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Mass Transfer (H2 O) in the Laboratory Environment
55 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Mass and Heat Transfer
56 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Mass and Heat Transfer
57 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Mass Transfer
dM
= Ka A(1 − FA )(Pa − Pu ) + Ks AFA (Ps − PL )
dt
58 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Mass Transfer
dM
= Ka A(1 − FA )(Pa − Pu ) + Ks AFA (Ps − PL )
dt
Atmosphere
Medium
!
"#
$ !
"#
$
dM
= Ka A(1 − FA )(Pa − Pu ) + Ks AFA (Ps − PL )
dt
59 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Mass Transfer
dM
= Ka A(1 − FA )(Pa − Pu ) + Ks AFA (Ps − PL )
dt
Atmosphere
Medium
!
"#
$ !
"#
$
dM
= Ka A(1 − FA )(Pa − Pu ) + Ks AFA (Ps − PL )
dt
What about the mass transfer coefficients?
60 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Mass Transfer Coefficients
1
1
∆x
1
=
+
+
K
Kb
DH2 O
Kl
61 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Mass Transfer Coefficients
1
1
∆x
1
=
+
+
K
Kb
DH2 O
Kl
1
1
∆x
=
+
K
Kb
DH2 O
Kb
DH2 O
∆x
is the film conductance of a layer of un-stirred air on shell
is the diffusion coefficient of shell divided by shell thickness
What about Mass Transfer Coefficients in the Soil?
62 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Top of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
M(1 − Fv )Cp
= Ha (1 − FA )A(Ta − Tu )
dt
63 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Top of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
M(1 − Fv )Cp
= Ha (1 − FA )A(Ta − Tu )
dt
Change of State of Water
!
"#
$
+ Hv Ka (1 − FA )A(Pa − Pu )
64 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Top of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
M(1 − Fv )Cp
= Ha (1 − FA )A(Ta − Tu )
dt
Change of State of Water
!
"#
$
+ Hv Ka (1 − FA )A(Pa − Pu )
Transfer within Egg
!
"#
$
hs Al
+
(TL − Tu )
R
65 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Top of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
M(1 − Fv )Cp
= Ha (1 − FA )A(Ta − Tu )
dt
Change of State of Water
!
"#
$
+ Hv Ka (1 − FA )A(Pa − Pu )
Transfer within Egg
!
"#
$
hs Al
+
(TL − Tu )
R
Metabolic Heat
!
"#
$
+ Hm (1 − Fv )
66 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Bottom of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
MFv Cp
= Hs FA A(Ts − TL )
dt
67 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Bottom of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
MFv Cp
= Hs FA A(Ts − TL )
dt
Change of State of Water
!
"#
$
+ Hv Ks FA A(Ps − PL )
68 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Bottom of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
MFv Cp
= Hs FA A(Ts − TL )
dt
Change of State of Water
!
"#
$
+ Hv Ks FA A(Ps − PL )
Transfer within Egg
!
"#
$
he Al
+
(Tu − Tl )
R
69 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Egg Heat Transfer-Bottom of Egg
Egg energy content
!
"#
$
Conduction
!
"#
$
dTu
MFv Cp
= Hs FA A(Ts − TL )
dt
Change of State of Water
!
"#
$
+ Hv Ks FA A(Ps − PL )
Transfer within Egg
!
"#
$
he Al
+
(Tu − Tl )
R
Metabolic Heat
! "# $
+
Hm F v
70 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Data
Solving the Model!
71 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Experimental Testing
72 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Experimental Testing
73 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Field Water Exchange
So What Happens with Egg Water Exchange in Field Nests?
74 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Snapping Turtle Eggs
75 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Average Mass Exchange All Clutches
76 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Field Water Potential at Nest Depth-Tensiometer
77 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Field Water Potential at Nest Depth-Tensiometer
78 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Characteristic Relation-Water Content and Potential (Ψ)
79 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Water Distribution
80 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Water Distribution
81 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Water Distribution
82 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Water Distribution
83 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Water Distribution
84 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Water Distribution
85 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Distribution of Water–Real Soil Bank
86 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Sea Turtle Beach, Nevatim Israel
87 / 88
Outline
Introduction
Temperature and Sex
Egg Water Exchange
Egg Heat and Mass Exchange
Egg Water Exchange in Soil
Sea Turtle Beach, Nevatim Israel
88 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
TSD
Appendix
89 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Gas Exchange
90 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Gas Exchange
91 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Gas Exchange
92 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Gas Exchange
93 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
94 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
95 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
96 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
97 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
98 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
99 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Heat Exchange
100 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
101 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
102 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
103 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
104 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
105 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
106 / 88
Appendix
Gas Exchange
Heat Balance
Water Balance
Clutch Water Exchange
107 / 88