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Age & GrowthReading: Chapter 9 (9.3)
Types of growthAging structuresBackcalculation methodsGrowth equationsParameter estimation
Individual processesImportant in understanding population dynamics
of fish• Fish are poikilotherms• Metabolic rate (and growth rate) affected by
temperatureImplications:
• Age and growth analyses• Timing of life history events (migration, spawning)Seasonal patterns are often the rule
Individual processes• Fish seek out preferred environments
(determined by temperature, dissolved oxygen, salinity)
Affects sampling and harvesting locations
• Early life history stages are less mobile; suffer high mortality in poor conditions
Evolution of high fecundity
2
Age & GrowthHow is growth
described?
– changes in length, width, or weight
– length is easiest
Age & Growth
Age & GrowthLength and age
Variety of methods to measure age• number and spacing of annual marks on a
part of the animal that is retained throughout its life
– scale, otolith, fin ray, vertebrae, spine, or shell
vertebrae
fin rays operclesotoliths
3
Age & GrowthRequirements of body structures to be used for
aging:1) structure must grow in constant proportion to
the size of the fish2) structure must exhibit easily-read periodic
marks that can related to time3) marks must be evident for all members of the
population4) marks must be constant across age groups and
across years
Ctenoid scale
4
Cycloid scale
regenerated scale
otolith-daily rings
5
Surf clam growth rings
Age & GrowthBackcalculation: Fraser-Lee Method
Proportional spacing of marks reflective of historical growth patterns
Fish size related to scale size by:
L = a + bS
where L = fish length, S = scale radius, and a = length at which scales start to form
Age & GrowthFraser-Lee method
Lengths at earlier ages can be backcalculated:
Li = length of fish at age iLc = current length of the fishSi = length of the scale at age iSc = current length of the scalea = correction factor (start of scale formation)
⎟⎟⎠
⎞⎜⎜⎝
⎛•−+=
=−−
c
ici
c
i
c
i
SSa)(LaL
SS
aLaL
6
Otolith size vs. body size
y = 20.1x - 9.4
0
200
400
600
800
1000
1200
1400
0 10 20 30 40 50 60 70Body size (cm)
Oto
lith
size
(mic
rons
)
Body size vs. Otolith sizey = 0.05x + 0.81
0
10
20
30
40
50
60
70
0 200 400 600 800 1000 1200 1400
Otolith size (microns)
Body
siz
e (c
m)
Data for spotted seatrout
Age & GrowthProportional methods
Structure proportional method:
Otolith size = 20.1(Body size) – 9.4
If: body sizecurrent = 50cmotolith sizecurrent = 900 micronsotolith sizeage1 = 180 microns
Then, use 4 steps to calculate body sizeage1
7
Age & GrowthProportional methods
Otolith size = 20.1(Body size) – 9.4(body sizecurrent = 50cm; otolith sizecurrent = 900 microns; otolith sizeage1 = 180 microns)
1. Calculate mean otolith size for a 50cm fish:
20.1(50) – 9.4 = 995.6 microns2. Calculate the ratio of observed otolith size to predicted mean otolith size:
900/995.6 = 0.9043. Adjust the observed otolith size at age 1 by this ratio to calculate the
expected otolith size for an age 1 sized fish:
180/0.904 = 199.14. Calculate the body size for which 199.1 is the expected otolith size:
199.1 = 20.1 (Body size) – 9.4 = 10.4cm
Age & GrowthFraser-Lee method
Lee’s phenomenon:tendency of back-calculated lengths from older fish to be smaller at early ages (age 1,2,etc.) than back-calculated lengths from younger fish in the populationWhy?.....Greater proportion of the larger fish in an age group dieother potential back-calculation errors (see p.195)
Age & GrowthLength & weight
Related by:W = a Lb
Above can be transformed:
lnW = lna + b*lnL
a and b can be derived from a ln/ln plot of weight as a function of length
8
Age & Growth:Length & weight
Data for a Lutjanus spp.
lnW = -11.11 + 3.04lnL
W = 1.5x10-6L3.04
Age & GrowthLength & weight
Related by:
W = aLb
Value of ‘a’ often used as an index of fish ‘condition’:
a = W / Lb
Not recommended; use ANCOVA instead to compare regressions
Age & GrowthGrowth
Expressed as the change in weight or length over time (ΔSize/Δt)Growth in fish often described by a logistic (or sigmoid) curve Same shape describes many biological functions in fish populations (individual and population growth, recruitment, size-selectivity of fisheries and predators, etc.)
9
Age
Len
gth
Hypothetical growth curve for fish
Age & GrowthCalculating growth
Absolute growth W2 - W1 / (t2 - t1)
Relative growth (W2 - W1) / W1
Instantaneous (lnW2 - lnW1) / (t2 - t1)growth
Age & GrowthGrowth curves
Describing size at age patterns:• underlying model must be biologically meaningful• von Bertalanffy model is most commonly applied• other models include linear, simple exponential,
Gompertz• parameters from von Bertalanffy model used in many
fishery yield models (to predict response to harvest)
10
Lt = length at time tL∞ = asymptotic length K = rate at which curve approaches L∞
t0 = hypothetical time when length equals zero
von Bertalanffy growth model
[ ]{ })( 01 ttKt eLL −−
∞ −=
von Bertalanffy growth model
von Bertalanffy growth model
11
Effects of variation in K
von Bertalanffy growth model
K and L∞ are species-specific (based on life history strategy)
von Bertalanffy growth model
So, what is t0?scaling factor related to juvenile growth
von Bertalanffy growth model
12
von Bertalanffy growth model
To convert from length to weight: -assume that W varies as a function of L3
[ ]{ }301 )( ttKt eWW −−
∞ −=
von Bertalanffy growth modelHow can we estimate the parameters?
Walford and Chapman plots, nonlinear models
[ ]{ })( 01 ttKt eLL −−
∞ −=
First step: Plot Lt+1 vs. Lt
von Bertalanffy growth modelWalford plot
Lt
L t+
1
1 to 1 line
13
From the plot of Lt+1 vs. Lt
1. Calculate the slope (b) = e-K (so, K= -lnb)
2. Y-intercept =
3. After re-arranging:
von Bertalanffy growth modelWalford plot
)( KeLa −∞ −= 1
baL−
=∞ 1
von Bertalanffy growth modelWalford plot
First plot Δ Lt which is (Lt+1 - Lt) vs. Lt
Then, the slope (b) = 1-e-K
Where the plotted line crosses the x-axis (x-intercept) = L∞
von Bertalanffy growth modelChapman plot
14
von Bertalanffy growth modelChapman method
Slope = 1 - e-K L∞
L t+1
-Lt
Lt
von Bertalanffy growth modelWalford and Chapman methods
t0 can then be estimated by substituting L∞ and K into the von Bertalanffy equation
∞
∞ −+=L
lLK
tt tln10
von Bertalanffy growth modelWalford and Chapman methods
Estimates of t0 will not be equally good for all lengthsGrowth curve will rarely pass thru originRemember, t0 is a scaling factor:– With negative t0 juveniles grow more quickly
than predicted growth for adults– With positive t0 juveniles grow more slowly than
predicted growth for adults
15
von Bertalanffygrowth model:
summary
Growth parameters from length-frequency plots
• Some species are difficult to age
• We can separate length-frequency distributions into cohorts and assign ages
• However, age-length relationships may not be valid if cohort separation is not clear
Length-frequencydistribution
Assigned age cohorts1
2
34 5
16
Short spawning season
fast growth rate
protracted spawning season
slow growth rate
+
+
Bhattacharya method
• Separates the length-frequency distribution into a series of normal distributions
• Identifies the youngest cohort and removes them from the distribution
• Approach is repeated
• Ages are assigned to each cohort and mean length at age calculated
overall length-frequency
first cohort identified
next cohort identified
17
ELEFAN method
Electronic length frequency analysisNo distributional assumptionsLength data are smoothed by taking running averages and best fitting growth curve is determinedMULTIFAN-a more objective alternative method
Data forChilean sea scallop