Stable Population Structure and Reproductive Value for Populations with Complex Life Cycles

Author(s): Hal Caswell

Reviewed work(s):
Source: Ecology, Vol. 63, No. 5 (Oct., 1982), pp. 1223-1231
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1938847 .
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Ecology, 63(5), 1982, pp. 1223-1231
? 1982 by the Ecological Society of America

STABLE POPULATION STRUCTURE AND REPRODUCTIVE

VALUE FOR POPULATIONS WITH COMPLEX
LIFE CYCLES'

HAL CASWELL2
Biological Sciences Group, University of Connecticut, Storrs,
Connecticut 06268 USA

Abstract. Classical demographic theory was developed to describe age-classified populations, in

which knowledge of an individual's age accurately specifies its demographic state. In many species
(e.g., plants, insects, fish) individuals of the same age may be in very different states; classical theory
is inappropriate for such populations. In this paper, I present general formulae for the stable stage
distribution (the analogue of the stable age distribution of classical demography) and the stage-specific
reproductive value distribution for populations with arbitrarily complex life histories. The results are
obtained by use of the z-transforms of a simple graphical representation of the life cycle. They yield
literal formulae, which can be used to draw qualitative conclusions about the general properties of
the life history. They will also provide quantitative results if numerical values for the parameters are
available. Since the selective pressures on life history traits can be expressed simply in terms of the
stable stage distribution and reproductive value, these results render complex life cycles amenable
to evolutionary analysis. The reproductive value formulae can sometimes be used to calculate optimal
schedules of stage-specific reproductive effort.
Key words: complex life cycles; eigenvalues; eigenvectors; life cycle graphs; life history theory;
matrix models; reproductive value; stable stage distribution; selective pressure; z-transforms.

INTRODUCTION stage-specific reproductive value distribution. These

Classical demographic methods focus on the dy- quantities, along with the long-term population growth
namics of age-structured populations (e.g., Keyfitz rate, are fundamental to demographic analysis and its
1968, 1977, Smith and Keyfitz 1977), using techniques evolutionary interpretation. Hubbell and Werner
originally developed for studies of human populations. (1979) have already generalized the characteristic
A classification of individuals by age in such a popu- equation, the solution to which gives the population
lation provides a reasonably accurate prediction of growth rate; this paper follows and extends their ap-
their demographic potential. In many species, how- proach. The theorems I will present yield formulae for
ever, individuals of the same age may be in very dif- the stable population structure and the reproductive
ferent demographic states. Age-classified demography value distribution in literal form, which is handy for
is incapable of dealing with such life cycles. Even in theoretical investigations. They will also provide nu-
human demography, it is sometimes necessary to clas- merical results if quantitative values are available for
sify individuals by other factors in addition to age the parameters.
(e.g., spatial location, parity, sex; Keyfitz 1968, Good- These results render complex life cycles subject to
man 1969, Rogers 1975). A number of authors have evolutionary analysis. The selective pressure on any
adapted the population projection matrix technique of life history parameter is given by the sensitivity of
Leslie (1945, 1948) to the complex life cycles of insects fitness to that parameter (Emlen 1970, Price 1970,
(Lefkovich 1965), herbaceous plants (Sarukhan and Crow and Nagylaki 1977), and this sensitivity can be
Gadgil 1974, Werner and Caswell 1977, Caswell and written down directly in terms of the stable population
Werner 1979, Bierzychudek 1981), trees (Usher 1969, structure and the reproductive value (Caswell 1978,
Hartshorn 1975, Enright and Ogden 1979), and fish Caswell and Werner 1978, Caswell and Hastings 1980).
(Horst 1977). Such analyses lack the theoretical basis In addition, optimal life history arguments can some-
provided for age-classified models by the work of Lot- times be phrased in terms of the maximization of re-
ka (1925), Fisher (1930), and Leslie (1945, 1948). The productive value (Schaffer 1974, 1979, Caswell 1980,
goal of this paper is to present two conspicuously 1982, Yodzis 1981).
missing parts of that theoretical basis: the general for- The results in this paper rely on a graphical repre-
mulae for the stable population structure and the sentation of the life cycle (the life cycle graph), on the
isomorphism between operations on such graphs and
the corresponding operations on matrices, and on the
1 Manuscriptreceived 15 April 1981;revised 25 November technique of z-transform analysis as applied to linear
1981;accepted 8 December 1981. discrete systems. In the next section, the formulation
2
Present address: Biology Department, Woods Hole
Oceanographic Institution, Woods Hole, Massachusetts of the basic demographic model is discussed. Subse-
02543 USA. quent sections analyze this model, examining the char-

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1224 HAL CASWELL Ecology, Vol. 63, No. 5

F~~ on this arc gives the number of individuals of stage i

at time t + 1 per individual of stage i at time t, that
3~~~ is:
04;? 0yD (a) ni(t + 1) = aijnj(t).
Graphs such as this have been used in life history
analysis before, notably by Lewis (1977), Gourley and
Lawrence (1977), and Hubbell and Werner (1979). My
usage differs slightly from that of Hubbell and Werner,
who include in their graphs arcs representing parti-
PP P2 P3 tions of stages within a single time interval and across
more than one time interval. Their graphs can always
G 3\> W b)
~~~~( be transformed into an equivalent graph of the form
defined here; the theorems to follow can be applied to
either type of graph.
The dynamics of the population are given by the
1 P2 3 P4
matrix difference equation

n(t + 1) =An(t), (1)

where n is a vector whose elements (ni) are the num-

P3 ~~F
2 graph of the matrix A (Coates 1959, Chen 1976).
FIG. 1. Life cycle graphs for three populations. (a) An
age-classified population, where Pi is the probability of sur-
viving from age i to age i + 1, and Fi is reproductive output
per individual of age i. (b) A size-classified population, where
Gi is the probability of growth from size-class i to i + 1, Pi
the probability of remaining in size-class i, and Fi the repro-
ductive output per individual of size-class i. (c) A hypothet-
ical plant population with both seed (F1) and vegetative (F2)
reproduction (see text for details).
bers of individuals in each of the different stages. A is
the population projection matrix, with aij equal to the
coefficient on the arc from nj to ni. (I will use the
symbol ni interchangeably to represent node i or the
number of individuals in the stage represented by node
i.) The life cycle graph is actually known as the Coates
Following these rules, the graph in Fig. la yields a
standard Leslie matrix, with the Fi in the first row and
the Pi on the subdiagonal. The graph in Fig. lb de-
scribes a size-classified population, in which an indi-
vidual in a given stage (size-class) may survive and
grow to the next size, survive and remain in its current
size, or reproduce. This graph yields a size-classified
matrix of the sort applied by Hartshorn (1975) and
Enright and Ogden (1979) to tree populations:

acteristic equation, the eigenvectors, and sensitivity P7 F2 F3 F4

analysis. Appendices contain discussions of the ana- A= G1 P2 ? ?
lytical technicalities and proofs of the theorems. o G2 P3 0
0 0 G3 P4
MODEL FORMULATION

As a starting point for analysis, choose a time scale
for the projection of the population, and construct a
life cycle graph (Fig. 1). The nodes of the graph rep-
resent demographically distinct classes within the pop-
ulation, i.e., they classify individuals by whatever
variables are necessary to determine their probabilities
of survival, reproduction, growth, movement, or other
demographic transitions. The form of the graph will
depend on the choice of the time interval (weeks,
months, years, etc.) over which the population is to
be projected. The nodes may be numbered in any con-
venient manner; I will choose n, to represent some
form of reproduction. A directed arc from node j to
node i represents a contribution to stage i at time t + 1
by individuals in stage j at time t. The coefficient aij
The possibility of remaining in the same category from
time t to t + 1, which is denied to age-classified pop-
ulations, appears as the diagonal elements of A and
the corresponding self-loops at each node of the graph.
Fig. Ic describes a hypothetical plant species with
both vegetative (F2) and seed (F1) reproduction. Seeds
germinate as seedlings (n1) which may develop in two
different sorts of microsites, say sunny (n3) and shady
(n2). By the following season, the plants are tall
enough that the microsite difference becomes irrele-
vant, and all individuals are lumped together in n4.
Plants of this stage may reproduce vegetatively, but
only in sunny microsites (F2). If they survive for
another year (n5) they reproduce by seed and die. The
resulting matrix is:

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October 1982 STABLE STRUCTURE AND REPRODUCTIVE VALUE
0 0 0 0 F1
P1 0 0 0 0
A P:3 0 0 F2 0
0 P2 P4 o o
o 0 0 P5 0
For the calculation of reproductive values, it is also
useful to consider the Coates graph of A', the trans-
pose of A. Since a'i aji, the transposed graph is
obtained by simply reversing the direction of every
arc in the graph. Rather than showing where individ-
uals of a given class may go, the transposed graph
shows where they may have come from.
The assumptions involved in this formulation of
population dynamics are those common to all matrix
projection models: linearity and time invariance. Hub-
bell and Werner (1979) distinguish life cycle graphs
from matrix projection models, on the grounds that
the graphs need not assume stable age distributions
within stages, and can deal with classes that are het-
erogeneous mixtures (e.g., dormant and nondormant
seeds). In fact, however, their (and my) analyses of
such graphs are completely isomorphic to analyses of
the corresponding matrices, and operate under exactly
the same assumptions. The graphs shown in their pa-
per are easily reduced to projection matrices with
identical properties.
The demographic analysis of Eq. I is straightfor-
ward. The largest of the eigenvalues (Xe)of A (denoted
here as X, without a subscript) is guaranteed to be real
and nonnegative by the Perron-Frobenius theorem,
and gives the asymptotic rate of population growth. I
assume, as is generally true, that X is strictly larger
than any of the other eigenvalues. This assumption
does not affect the formulae to be derived, only the
relevance of the eigenvalue. The right eignevector w
corresponding to X is defined by
AW = Xw
and gives the stable stage distribution, to which the
population will ultimately converge. The left eigen-
vector v is defined by
V'A = Xv',
and gives the stage-specific reproductive value distri-
bution, i.e., the value of the contribution of each stage
to long-term population growth. It is the discrete
equivalent of Fisher's (1930) formula in the special
case of age-classified populations; for a proof in gen-
eral matrix terms, see Goodman (1967) or Caswell and
Werner (1978). The evolutionary analysis of life his-
tories turns on the use of X as a measure of mean
fitness. Under slow selection in the absence of com-
plex epistatic interactions, the selective pressure on
a quantitative trait x is given by dX/dx (Emlen 1970).
For life history traits expressed as entries in the matrix
A, the corresponding selective pressures can be writ-
ten down directly in terms of w and v (Caswell 1978).
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1225
Thus knowledge of w and v provides the key to eval-
uating the evolutionary importance of the various ele-
ments in the life history.
ANALYSIS
The derivations of w and v begin with the z-trans-
form of the life cycle graph. This transform is the dis-
crete analogue of the Laplace transform for differential
equations (Jury 1964, Schwarz and Friedland 1965).
To obtain the z-transform of the life cycle graph as
defined in this paper, simply replace the coefficient aij
on the arc from nj to ni by aijX-1. The exponent of
-1 corresponds to the single time unit lag in the
expression ni(t + 1) = aijnj(t). If there are arcs in the
graph which correspond to other than unit delays (in
which case the graph will not correspond as directly
to the matrix), the (negative) exponent on X corre-
sponds to the length of the lag (Hubbell and Werner
1979).
There is a notational dilemma here. In the engi-
neering literature, the transformation would be written
aiiz-1; hence the name. However, the symbol used will
shortly be equated with the eigenvalue of A, custom-
arily denoted by X. The choices seem to be: change
the name of the transform, change the symbol for the
eigenvalues, or be inconsistent. I have chosen the last
alternative.
The characteristic equation
The characteristic equation, the solutions of which
are the Xi, can be written down directly from the z-
transformed graph (Hubbell and Werner 1979), using
Mason's expansion of the graph determinant (Mason
and Zimmerman 1960, Huggins and Entwisle 1968, see
Appendix A). (Levins' [1974, 1975] technique of loop
analysis is also based on Mason's formula, but differs
in several respects from the approach here. He applies
the formula to the untransformed graph of a continu-
(2)
ous, rather than a discrete system. His main concerns
are stability [evaluated using the Routh-Hurwitz
criterion] and the sensitivity of equilibrium points
[evaluated using Cramer's rule]. Eigenvectors do not
enter in.)
(3)
Let a loop be defined as a closed path in the graph,
traversed in the direction of the arrows and not in-
cluding any node more than once. The transmission
L of a loop is the product of the coefficients around
the loop (with a slight complication introduced to deal
with self-loops; see below). The determinant of the
graph is given by
I - Li + a* LiLj - Li
LjLLk +.... (4)
i ij ijk
The first summation is the sum of the transmissions of
all the loops in the graph. The subsequent summa-
tions, marked with asterisks, add the products of all
pairs, triplets, etc., of disjoint loops, that is, loops that
share no nodes.
1226 HAL CASWELL Ecology, Vol. 63, No. 5

The characteristic equation of the matrix A is ob- w= 1, (6a)

tained by setting the determinant of the z-transformed
graph equal to zero (Appendix A). Given all those X-1
terms in the z-transformed graph, the characteristic
equation is a polynomial in X-1. Solution of this poly-
nomial yields the eigenvalues of A; Hubbell and Wer-
ner (1979) give detailed instructions and examples.
Notice that the loops and their relationships are un-
changed when the graph is transposed. Thus, the
transposed matrix has the same characteristic equa-
tion and eigenvalues.
The presence of self-loops in a graph requires a de-
cision. They may either be counted as loops in the
determinant formula, or they may be absorbed into
the calculation of the transmissions. The formulae re-
sulting from the latter choice are much simpler (since
self-loops on different nodes are always disjoint), and
will be used throughout this paper. The presence of a
self-loop of strength ajjX-1at node nj divides the trans-
mission of all incoming arcs by (1 - ajjX-1). Once this
division is performed, the self-loop is eliminated from
the graph. For details see Mason and Zimmerman
(1960) or Hubbell and Werner (1979).
The stable stage distribution
To calculate the right eigenvector w, which gives
the stable stage distribution, define the following paths
and loops in the z-transformed life cycle graph:
Ti,= the transmission of the ith directed path
from n1 to n,. The subscripts denote the
beginning and the end of the path; the su-
perscript denotes which of the possibly nu-
merous paths between the two points is
under consideration,
Lj(n,) = the transmission of the jth loop which is
disjoint with node n1, and
Lj(T1i) = the transmission of the jth loop, which is
disjoint with the path 71x,
and
Wx = E Tr, for x > 1. (6b)
In this case, there is a simple recursive formula for
wX, with w1 = 1, and
WX= TjXwj. (7)
The summation is taken over all stages j which feed
individuals directly to nx, that is, all stages which are
only a single stage "upstream" from nx.
These formulae are most easily understood in the
form of Eq. 6. The elements of w appear there scaled
so that w1 = 1. To find the number of stage x individ-
uals per stage 1 individual, one simply adds the trans-
missions of all the paths by which stage I can contrib-
ute to stage x. This interpretation is intuitive, but
could not be applied to the untransformed graph be-
cause of the time taken to develop from n1 to nx, and
the changes in population size occurring during that
time if X 5 1. In effect, the act of taking the z-trans-
form of the graph assures that the delays involved in
the different pathways, and the effect of the rate of
growth of the population, are properly taken into ac-
count. The recursive form (Eq. 7) is similar in concept;
the abundance of ni in the stable population is ob-
tained here relative to the abundances of the stages
which contribute to it directly.
Examples.-The graphs in Fig. la, b satisfy the con-
ditions for Eqs. 6 and 7, so the stable stage distribu-
tions and the recursive formulae can be written down
directly. For Fig. la,
WI = 1
W2 = P1X-l
W3 = P1P2 X-2
W4 = P1P2P3X-3

Theorem 1.-The elements of the stable stage dis- WX = Px-iX-1Wxi,

tribution vector w are given by
and for Fig. lb,
WI = 1- Lj(n) + Lj(nl)Lk(nl) -. . ., (5a)
j j,k WI = I
W2 = G1/(X - P2)
and
W3 = G1G2/(X - P2)(AX- P3)
W4 = - P3)(AX
GlG2G3/(X - P2)(AX -P4)
Tx41 Lj(Tix)
w, =E i j
WX = GX_1W1/(X - PX).
+ E* Lj(Tix)Lk(Tx)- . .]. (5b)
j, k The formulae obtained from the graph of Fig. la agree
The summations marked by asterisks are taken over with those obtained by standard methods from the
pairs, triplets, etc., of mutually disjoint loops. (See Leslie matrix (e.g., Pollard 1973). Note that the re-
proofs, Appendix B.) cursive formula for the size-structured model differs
Before discussing this theorem, it will be helpful to from that for the age-structured model, only in the
examine a couple of special cases. If all reproductive appearance of (X - Px) in place of X in the denomi-
paths feed back through a single stage (call it stage 1 nator.
for convenience), there will be no disjoint loops in Eq. The presence in Fig. lc of the loop P412, which does
5a, b, and the stable stage distribution reduces to not feed back through n1, requires the use of the com-

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October 1982 STABLE STRUCTURE AND REPRODUCTIVE VALUE 1227

plete formulae (Eq. 5a, b). The resulting formula for calculated with self-loops dividing outgoing rather
W is: than incoming arcs. Thus
WI = I -P4F2X-2 T
Ti (-all)
W2 = P1X-1(l -P4F2X-l) -
(X a ~X~1-
W3 = P3XcI + P1P2F2X-3
W4 = PiP2X-2 + P3P4X-2 Theorem 2.-The elements of the left eigenvector
W5 = PI P2P5X-3 + P3P4P5X-3. v are given by

The presence of the loop (P4F2) complicates things in VI= 1 - Lj(n1) + E Lj(nl)Lk(nl)-. . ., (8a)
j j,k
two ways. First, w1 no longer equals 1, so the other
values cannot be interpreted as individuals of stage x and
per individual of stage 1. If desired, all the values X= X - Lj(ti )
i
could be divided by (1 - P4F2X-2) to achieve this.
Second, the existence of this loop renders the recur- + A*Lj(txiJLk(txi1)- (8b)
sive formula for wX incorrect in general. j, k

The parameter X in these formulae is the eigenvalue The summations marked with asterisks are taken over
of A, obtained by solving the characteristic equation. pairs, triplets, etc., of disjoint loops. (See proofs, Ap-
If numerical values for all the parameters were avail- pendix B.)
able, the characteristic equation could be solved for Again, the formulae are considerably simplified if all
X, and the resulting value used to compute numerical the loops in the graph pass through n1, in which case
values for w. In most such situations, however, it will the reproductive values are
be easier to obtain the eigenvalues and eigenvectors
v= 1, (9a)
directly by computer. More interest is likely to attach
to the interpretation of the literal expressions. For ex- and
ample, the stable age distribution is always a strictly
Vx = T, for x > 1. (9b)
decreasing function, as long as X > 1, i.e., there are
more individuals in each age-class than in the suc-
ceeding one. It is easy to see that this is not true for In this case, there is also a recursive formula for the
the stable size distribution obtained from Fig. lb. In elements of v. Let FX be the coefficient on the path (if
fact, if Gj > (X - Pj+,), then wj+,1> Wj. If the proba- any) directly from nX back to n1. Then,
bility of growing into stage j + 1 is large, and the prob-
ability of dying or growing out of it is small, then there VX= Fx +TXiVi (10)
will be a hump in the stable size distribution, with -axxi

individuals stacking up in stage j + 1. Such humps in

size distributions are well known in fish and tree pop- where the summation is taken over all the stages to
ulations. These results show that they may be part of which stage x is directly connected.
the stable population structure, rather than transient The contrast of Eq. 9 with Eq. 6 is revealing. While
phenomena resulting from perturbations to the stable the elements of w are transmissions from n1 to nx, the
structure. elements of v are transmissions from nX back to n1.
Although demography focuses primarily on the To obtain the reproductive value of a given stage, one
maximal eigenvalue of A, the formula for w (and v simply adds all its contributions to reproduction, with
below) applies equally well to the eigenvectors cor- the z-transform automatically weighting the different
responding to any of the other eigenvalues (with ap- reproductive pathways to include the effects of delays
propriate substitution for X). Since the subsidiary ei- and of population growth. If there are loops in the
genvalues of A help to determine the transient graph which do not pass through stage 1, the formulae
behavior of population, analysis of these eigenvectors get more complex, but the principle is the same.
may eventually be of interest. The recursive formula (Eq. 10) was first used in the
age-classified case by Williams (1966), who partitioned
Reproductive value the reproductive value at age x into current reproduc-
The reproductive value vector, v, is the left eigen- tion (Fx) and what he termed "residual reproductive
vector of A corresponding to X. In addition to the def- value." In the age-classified case, the residual repro-
initions for Theorem 1, define ductive value is the probability of survival to the next
age times the reproductive value at that age. In the
TXh= the transposed transmission of the ith path from general case, reproductive value is partitioned into
nx to n1 (i.e., the ith reproductive path). current reproduction (including the possibility of sur-
viving to reproduce again in the same stage), plus a
Because of the reversal of the arcs associated with residual reproductive value which sums the probabil-
the transposed graph, the transposed transmission is ity of movement to perhaps several "next" stages,

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1228 HAL CASWELL Ecology, Vol. 63, No. 5

times their reproductive values. In practice, this for- Goodman 1971, Keyfitz 1971). A general solution of
mula is easy to use backwards. One begins by calcu- this problem, valid for any matrix model, leads to the
lating the reproductive values of all stages which con- simple formula (Caswell 1979):
tribute only to n1 and then works back to the stages
aX/0aij v:iwj/(v,w),
= (11)
contributing to those stages, and so on.
Examples.-The reproductive values for the life where ( denotes the scalar product.
cycle in Fig. Ia are Knowledge of v and w, together with Eq. 11, suf-
VI = I
fices to determine the importance to fitness of any de-
v2 = F2X-' + P2F3X-2 + P2P3F4 - 3 mographic parameter. If numerical values for A are
V3 = P3X1- + P3F4X-2 available, the sensitivities can be calculated quantita-
V4 = F4X-I tively. Often, however, qualitative results based on
V, = FxX-1 + PxX-Xvx+1.
literal formulae are more useful than special cases
based on quantitative data. In such calculations, the
For the size-classified graph (Fig. lb), v is scalar product term in the demoninator of Eq. 11 is
VI = I
not stage dependent, so it is usually easier to think in
terms of relative sensitivities, and use just the numer-
_ F2 G2F3
+ ator.
(XA-P2) (X-P2)(X-P3) As an example, consider the problem of senescence
+ G2G3F4
in age- and size-classified populations. Most species
(A-P2A- P3)(AX-P4) exhibit senescence, and several evolutionary expla-
V3 - F3 + G3F4 nations of it have been attempted (Medawar 1952,
(X-P3) (X-P3)(AX-P4) Williams 1957, Hamilton 1966). In age-structured pop-
F4
ulations, the relevant sensitivities are
v=
(X - P4)
i~aAF, = vwx,' (12a)

vx= Fx + Gxvx+l and

X
(A-PX) (X-PX)
ax/OPx = Vx+iWx. (12b)
Note that the formula for the age-specific reproductive
value agrees with the standard formula due to Fisher The first of these quantities always declines with
(1930, see Pollard 1973 for the discrete time version). age, at least when X D 1. The second is also mono-
The presence of the loop (P4F2) in Fig. lc makes tonically decreasing in a non-senescing population (the
things slightly more complex, but direct application of appropriate initial condition for examining the evolu-
Eq. 8 yields: tion of senescence). Thus genes with small beneficial
=
effects early in life and large deleterious effects late in
V I - P4F2X-2
life may have a net positive effect on fitness, and be
V2 = P2P5F1X-3
selected; the result is senescence. A non-senescing
V3 = P4P5F1X-3
population is, in effect, evolutionarily unstable. The
V4 = P5F1X-2
- situation in size-classified populations is fundamental-
V5 = FI-'(I lP4F2X-2).
ly different. We must now consider:
Note that the recursive formula (Eq. 14) no longer
holds. a~a/Fx= V1wx, (13a)

Sensitivity analysis axlaPX= VxWx, (13b)

A fundamental problem in life history theory is to
determine the contribution of the various parts of a
life cycle to fitness. (The choice of a measure of fitness
is probably more fundamental; here I am following the
standard practice and using X as such a measure. This
choice tacitly ignores the problems of variable envi-
ronments, diploid genetics, and nonstable stage dis-
tributions.) What is needed is a way to trace the effect
of a change in the life history through the life cycle
graph, the associated matrix, and the characteristic
equation, arriving finally at a resulting change in X. In
age-structured models, the characteristic equation is
well known, and several authors have used it to derive
the sensitivity of X to changes in age-specific param-
eters (Hamilton 1966, Demetrius 1969, Emlem 1970,
and
Wa/Gx = vx+1wx- (13c)
Since neither wX nor vX need be decreasing functions
of size, effects at larger sizes may be more important
than effects at smaller sizes. In such circumstances,
the selective pressure favoring senescence will be
weaker or nonexistent. This was suggested long ago
by Bidder (1925, 1932) for fish with indeterminate
growth, but based on a group-selection argument.
However, it is interesting to note that the cases in
which senescence seems least pronounced or even ab-
sent (Comfort 1964, Harper 1977) are in organisms
whose plastic growth makes them candidates for a
size-classified demography. Note that this result re-

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October 1982 STABLE STRUCTURE AND REPRODUCTIVE VALUE
quires the size-structured life cycle. Even if an age-
classified population has continually increasing repro-
ductive output (and thus reproductive value), it is still
vulnerable to the evolution of senescence (Hamilton
1966).
The eigenvector formulae can be used to examine
other kinds of tradeoffs as well. Why, for example,
does stage 4 in Fig. lc not indulge in sexual as well as
vegetative reproduction? There is evidence that the
two modes of reproduction are antagonistic (e.g.,
Abrahamson 1980), so one hypothesis is that the cost
of adding seed reproduction is greater than its benefits.
Let F3 represent seed production by n4, and let the
cost be given by dF2/dF3 = - c. The condition for an
increase in F3 (from its value of zero) to be favored
is:
dX= [aF3 -c
(9
jdF3 > O.
So if the hypothesis is true, then
axa/F3 < C.
AA/8F2
Evaluating the partials using Eq. 11, this condition
reduces to
V- I -P4F2A-2 < C.
V'3 P4P5F1X3
Thus, the hypothesis that F3 = 0 because of its cost
might be tested by looking for situations where the
ratio in Eq. 14 becomes large, i.e., where P5 or F1
become very small. In such populations, the evolution
of seed production by n4 might be favored.
Finally, note that reproductive value also plays an
important role in Schaffer's (1974, 1979; see also Wil-
liams 1966, Taylor et al. 1974) analysis of life history
optimization in terms of reproductive effort. He
showed (cf. Yodzis 1981) that, in an age-classified pop-
ulation, the schedule of reproductive effort that max-
imizes X also maximizes reproductive value at each
age. This important result can be extended, under
some conditions, to stage-classified populations (Cas-
well 1982). General expressions for the form of v may,
then, allow optimal reproductive effort calculations to
be made for stage-classified populations.
ACKNOWLEDGMENTS
This research has been supported by the National Science
Foundation through grant DEB/OCE 76-19278. I thank Dan-
iel Goodman and an anonymous reviewer for helpful com-
ments on an earlier draft.
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APPENDIX A
Mason's formula and Coates' graph
The connection between the determinant of the z-trans-
formed life cycle graph and the characteristic equation of the
population projection matrix requires some clarification.
There are two common ways of associating a graph with a
matrix, one called a signal flow graph (Mason 1953, 1956) and
the other a flow graph (Coates 1959). To avoid confusion, I
follow Chen (1976) and refer to them as Mason graphs and
Coates graphs. There are formulae for the evaluation of de-
terminants, etc., for both types of graph, but while they lead
to the same answers, they involve different operations. The
life cycle graph is the Coates graph of A, but the characteristic
equation is obtained by applying Mason's formula for the
determinant. Why does this work?
Mason graphs and Coates graphs are related by
GC(A) = Gm(A - 1), (Al)
where G, and Gm stand for Coates and Mason graphs. Con-
sider the z-transformed Coates graph
G, (Z'-'A) = Gm(Z'-'A - I). (A2)
That is, the z-transformed life cycle graph is equivalent to
the Mason graph of (z-1A - i). Application of Mason's ex-
pansion for the determinant to this graph yields
det(z-1A - i) = z-8det(A - ZI), (A3)
where s is the order of the matrix A. Thus when Eq. A3 is
set equal to zero, the result is the characteristic equation,
and the values of z which satisfy it are the eigenvalues of A.
APPENDIX B
Proofs
Although the eigenvector formulae are based on a rela-
tively straightforward extension of Mason's graph theory, I
have not been able to find them in the engineering literature.
The proof presented here is original; I would welcome ref-
erences to earlier proofs if they exist.
Theorem I.-Let B = (A - Xi) be the characteristic matrix
of A (i.e., the matrix whose determinant appears in the char-
acteristic equation). The right eigenvector w is the solution
to BW = 0. B, however, is singular (since the eigenvalues are
obtained by setting its determinant to zero), and thus cannot
be inverted to solve for w. A solution for w can be written
in terms of the cofactors of B. Let Bi, be the (ix) cofactor
of B, that is, the determinant of the matrix formed by deleting
row i and column x from B. Then wX = Bix for any i (Chiang
1968). The formula for w in Theorem 1 results from applying
this result with i = 1.
The cofactor B~xis equal to the determinant of the matrix
formed by replacing row I and column x of B with zeros, and
placing a I in the (1 ,x) position. The graph manipulation cor-
responding to this alteration in the matrix is as follows: (1)
absorb all the self-loops, (2) remove all arcs leaving nx and
entering n1, (3) add an arc with coefficient I from nx to n1,
and (4) add self-loops of strength I to n1 and nx. (The final
manipulation corresponds to eliminating row I and column
x of the identity matrix subtracted from Z-1A in Eq. A3.)
As a result of this manipulation, all loops passing through
n1 or nx have been broken, save those passing from n1 to nx
October 1982 STABLE STRUCTURE AND REPRODUCTIVE VALUE 1231

and back, each of which has a loop transmission equal to
TiX. There are two self-loops of strength I (or one of strength
2 when x = 1). Any other loops remaining in the graph are
disjoint with all the Tir, and with the unit self-loops. Apply-
ing Mason's determinant formula to this graph leads to the
expression in Theorem I (see Theorem 3.8 of Chen 1976).
Theorem 2.-The left eignevector v is the right eigenvector
of A', and the graph of A' iS obtained by reversing the direc-
tions of all the arrows in the graph of A. Theorem 2 is ob-
tained by applying Theorem I to the transposed graph, and
then expressing the results in terms of the original graph. The
loops and the disjointness relations between them are unaf-
fected by the transpositon. A directed path from n1 to n, in
the transposed graph is a path from n, to n1 in the original
graph. The transmissions of the two paths differ only because
of the effect of self-loops. In the transmission from n, to n,
in the transformed graph, self-loops divide incoming arcs. If
the transmission is expressed in terms of the path from n, to
n1 in the original graph, self-loops must divide outgoing arcs.
This change affects only the endpoints of the path. Theorem
2 follows immediately from applying Theorem I to the trans-
posed graph.

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