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growth matrices

Ramón Filgueira, Laura G. Peteiro, Uxı́o Labarta

and Marı́a José Fernández-Reiriz

C.S.I.C.—Instituto de Investigaciones Marinas, C/Eduardo Cabello 6, 36208 Vigo, Spain

The increase in size of individuals in a population implies an Fréchette & Lefaivre (1990) established a method to dis-

increment of their space and food requirements. These factors tinguish between both causes of competition – space or food –

when limiting can cause intraspeciﬁc competition, resulting in in the relationship between the maximum biomass that can be

a decrease of the growth rate and an increase in the mortality supported by a population and its density (self-thinning). The

rate (Alunno-Bruscia et al., 2000). The self-thinning process method is based on the equations described above, ascribing to

describes the negative relationship between body size and the space-limiting factor (spatial self-thinning, SST) the

population density when growth results in mortality through relation m ¼ kN 23/2 (or B ¼ kN 21/2) and to the food-limiting

intraspeciﬁc competition (Westoby, 1984). Self-thinning can be factor (food self-thinning, FST) the relation m ¼ kN 24/3 (or

studied by means of successive sampling of the growth of B ¼ kN 21/3).

even-aged individuals that are maintained at different densities In the present study we analysed a dataset of dry weights

(see Fig. 1 in Alunno-Bruscia et al., 2000). The following and densities of Mytilus galloprovincialis cultivated in the Rı́a de

equation describes the self-thinning rule (Westoby, 1984): Ares-Betanzos (NW Spain). The data came from the regular

sampling carried out by our research group for various other

studies. At each sampling, two replicates of 50 cm of rope were

B ¼ kN 1=2 ; ð1Þ

scraped free of mussels. The mussels were counted, weighted,

measured and classiﬁed into length classes. The data were

where B is the total biomass, k the intercept of the relationship taken during the years 2004 (n ¼ 111), 2005 (n ¼ 149) and at

and N the population density. The last expression is obtained the beginning of 2006 (n ¼ 8). The use of data sampled across

from the mean morphometric relationships between the sub- years allowed us to consider the assumption of constant energy

stratum area occupied by the individual (S), length of the indi- ﬂow less likely violated because ﬂuctuations in food abundance

vidual (l) and its weight (m). Assuming isometric growth (S a l 2 over years should only cause increased variance around the

and m a l 3) and an inverse relationship between density and self-thinning line (Steingrı́msson & Grant, 1999). Mussels were

area occupied (N a S 21), the following equation is obtained: cultivated on ropes at high densities, which presented an ideal

situation to study the self-thinning processes because this

m ¼ kN 3=2 ; ð2Þ method involves mortality by intraspeciﬁc competition. The

diameters of 25 commercial ropes were measured to calculate

which given the relationship between population biomass and the available area for mussel attachment. Ropes were hung

individual average weight (B ¼ Nm) can be expressed as from rafts following the technology used commonly in the

Equation 1. Galician Rı́as, which includes three phases: seed settlement

Although this theory is based on studies focused on plants, it (initial), early thinning-out and thinning-out. In the two last

can be applied to both sessile and mobile animal populations phases, the mussels are detached from the ropes and replaced

(Hughes & Grifﬁths, 1988; Steingrı́msson & Grant, 1999). in culture at lower densities. The reduction of the density

Damuth (1981) observed in primary consumers an exponent of caused by this manipulation alters the natural process of self-

24/3 between average weight and density and suggested that thinning and invokes mitigation in the mortality by intra-

this can be related to the metabolic requirements of indivi- speciﬁc competition. The subsequent inclusion of values in

duals. In this way, Begon, Firbank & Wall (1986) stated a which the mortality by intraspeciﬁc competition has not yet

theoretical relationship between the self-thinning process and begun will ﬂatten the estimated slope of the self-thinning ﬁt

the population metabolic rate (MR), which is related to the (Zhang et al., 2005). Several studies have suggested corrective

individual average weight (MR a Nm 3/4) and environmental techniques that include the removal of these data points

conditions (MR a F, where F is the total amount of food con- (Westoby, 1984) or the use of different regression models that

sumed by the population). Assuming a constant consumption allow a good ﬁt without the omission of data points (see review

of food by the population (Fcte), in stable environmental con- by Zhang et al., 2005). These last authors compared several

ditions the following relationship can be obtained: regression models [quantile regression, deterministic frontier

function and stochastic frontier function (SFF)] that have the

potential to establish an upper limiting boundary line above

m ¼ kN 4=3 ; ð3Þ all plots for the maximum size –density relationship, without

subjectively selecting a subset of data points based on pre-

which given the relationship between population biomass and deﬁned criteria. Zhang et al. (2005) concluded that the SFF

individual average weight (B ¼ Nm) can be expressed as: was the most suitable model given that this method can easily

yield statistical inference on the model coefﬁcients.

In the present study, the mathematical expression that

B ¼ kN 1=3 : ð4Þ

describes the self-thinning (m ¼ kN g) was calculated by means

of the logarithmic linear transformation (log m ¼ log

Correspondence: M.J. Fernández-Reiriz; e-mail: mjreiriz@iim.csic.es k þ g log N) following two regression models: reduced major

# The Author 2008. Published by Oxford University Press on behalf of The Malacological Society of London, all rights reserved.

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RESEARCH NOTE

layered structure of an intertidal mussel community is taken

into account. In this model, the individual density (N) is con-

sidered inversely proportional to the area projected on the sub-

stratum (S) and directly proportional to the number of layers

(L), i.e. NaL S 21. Therefore, the B–N–L tridimensional

model that deﬁnes the self-thinning can be expressed in its log-

arithmic form as (Guiñez & Castilla, 1999):

and density (N), the B–N–L tridimensional model could be

Figure 1. Dry weight of individual mussels vs culture density, and reduced to a B–N bidimensional model, although this change

linear ﬁts performed by RMA and SFF. ANCOVA following Zar alters the exponent of the model and, therefore, it could not be

(1984) was performed to compare the observed exponents with compared with the theoretical exponents for space- and

theoretical values for FST and SST (24/3 and 23/2, respectively). food-limiting factors. Guiñez & Castilla (2001) suggested a

new tridimensional model that allows comparison of the

empirical exponent with the theoretical exponents for space-

axis (RMA) and SFF (Zhang et al., 2005). The regression and food-limiting factors. This model introduces the concept of

models were carried out using the statistical package SPSS effective density (Ne), which is deﬁned as the expected density

14.0 and Frontier 4.1c (Coelli, 1996), respectively. The results if individuals within a sample occurred as a monolayer. The

shown in Figure 1 highlight the importance of the regression log L term of the tridimensional model is then removed and

model used for ﬁtting the self-thinning equation. The RMA the model can be expressed as:

regression results in a slope of 21.58 + 0.106 (mean + 95%

conﬁdence interval), which is statistically similar to the theor- log m ¼ log k þ g log Ne : ð6Þ

etical slope of the space limitation relationship (SST;

ANCOVA: t ¼ 1.481, P ¼ 0.140, n ¼ 268), whereas the SFF The application of this model requires knowledge of the area

regression results in a slope of 21.16 + 0.102, which is statisti- occupied by each individual. In this way, assuming that the

cally different from the theoretical values for FST (ANCOVA: maximum length of an individual is perpendicular to the sub-

t ¼ 3.213, P , 0.005, n ¼ 268) and space limitation (SST; stratum, Guiñez & Castilla (1999) estimated the area projected

ANCOVA: t ¼ 6.476, P , 0.001, n ¼ 268). Although the RMA onto the substratum by multiplying the maximum width by

has yielded an exponent similar to the theoretical value, the the maximum height; that is, assuming that the mussel is con-

handling of cultivation ropes, with a consequent reduction in tained in a parallelepiped. The application of this model to the

mussel density, introduces data points for which mortality by present study dataset results in a reduction of the exponent

intraspeciﬁc competition has not occurred. So, caution must be from 21.16 + 0.102 to 21.62 + 0.024 (Figs 1 and 2A, respect-

taken in interpreting the meaning of the self-thinning exponent ively), which is statistically different from the theoretical values

estimated by means of the RMA regression model. Therefore, for FST (24/3; ANCOVA: t ¼ 24.167, P , 0.001, n ¼ 268)

the comparison between theoretical values and the self- and SST (23/2; ANCOVA: t ¼ 10.000, P , 0.001, n ¼ 268).

thinning exponent calculated by means of the SFF (–1.16 + The exponent reduction is caused by the use of effective

0.102) indicates that the population’s regulation is caused by density, which corrects the underestimation of the bidimen-

neither space nor food limiting factors. sional model in which the multilayered structure is not con-

However, distinguishing between space or food limiting sidered (Guiñez, Petraitis & Castilla, 2005).

factors by means of the value of the exponent is a difﬁcult task Mussels are gregarious animals that form complex matrices

because it is possible that both limiting factors exert a density- with several overlapped layers (Fig. 3), in which the indivi-

dependent effect on sessile populations (Fréchette, Aitken & duals that are placed in the external layers are overlapped with

Pagé, 1992), and the exponent could show deviations from the the internal layers, occupying the empty space of the internal

theoretical value (Alunno-Bruscia et al., 2000). In an analogous layers. Therefore, calculating the occupied area by means of

way, wide variability has been observed in the exponent of the the parallelepiped projection, deﬁned as the exclusive indivi-

relationship between MR and weight (Latto, 1994), which dual space occupied per mussel, could overestimate the real

would modify the theoretical value of the FST given the area occupied by the mussels because the overlapping is not

dependence of MR and self-thinning established by Begon considered. An approach more representative of mussel mor-

et al. (1986). In the case of a space factor, the assumptions of phology could yield a more realistic value of the occupied area

the classic model are: (1) isometric growth, (2) a complete per individual. Given the predominant position of the mussel,

occupation of the sampled area and (3) an inverse relationship with the maximum length of the individual perpendicular to

between the sampled area and the individual density (NaS 21; the substratum, the apical projection onto the substratum

Westoby, 1984). However, growth usually is allometric instead could be a better approach to determine the occupied area. In

of isometric causing a deviation from the theoretical exponent this way, the projected area occupied by the population will be

(Westoby, 1984). In addition, the multilayered structure more realistic as the population packing increases. An accurate

formed by the mussel on the substratum violates the third tool to determine complex areas is image analysis. In the

assumption and therefore alters the theoretical exponent present study, apical photographs (Nikon Coolpix 4500) of 50

because the available area is underestimated (Hughes & individuals (from 15 to 90 mm length) were taken to deter-

Grifﬁths, 1988). These deviations have been included in newer mine the projected area onto the substratum by means of

models (Fréchette & Lefaivre, 1990; Guiñez & Castilla, 1999) ImageJ 1.37v. The allometric antero-posterior axis length vs

of mussel growth that modify classical models to introduce the projected area (projected area ¼ 0.8 1023 length2.22, N ¼

effect of allometric growth, multilayered structure and/or the 50, r 2 ¼ 0.99, P , 0.001) was applied to the frequency distri-

effect of the roughness of the substratum. Guiñez & Castilla butions to calculate the occupied area of the population. The

416

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RESEARCH NOTE

Figure 2. Dry weight of individual mussels vs culture density, and linear ﬁts performed by RMA, where N is the effective density calculated from

the projected area of the parallelepiped (A), and where N is the effective density calculated from the image analysis (B). In both cases, only RMA

is shown because the SFF estimates identical parameters and the likelihood value is less than that obtained using RMA. ANCOVA following Zar

(1984) was performed to compare the exponents observed in the present study with theoretical values for FST and SST (24/3 and 23/2,

respectively).

viduals by the occupied area of the population. Figure 2B

shows the average weight vs density, expressed as number of We thank PROINSA mussel farm and their employees,

individuals per effective area calculated by means of image especially H. Regueiro, M. Garcı́a, C. Brea and

analysis. The observed slope value (21.32 + 0.027) is statisti- O. Fernández-Rosende for technical assistance. We wish to

cally similar to the theoretical exponent of 24/3 (ANCOVA: thank A. Ayala for helping us with Figure 3.

t ¼ 1.111, P ¼ 0.268, n ¼ 268), which would indicate that the

population growth is limited by available food (Fréchette & FUNDING

Lefaivre, 1990). The application of image analysis techniques

allow the calculation of a more realistic value of occupied area, This study was supported by the contract-project PROINSA,

which in the particular case of Mytilus galloprovincialis conﬁrms Code CSIC 20061089, Xunta de Galicia PGIDIT06RMA018E.

that in densities commonly used in commercial cultivation on

rafts, self-thinning is probably regulated by the food-limiting

factor. REFERENCES

The arrangement of mussels in complex matrices makes it ALUNNO-BRUSCIA, M., PETRAITIS, P.S., BOURGET, E. &

difﬁcult to apply classical self-thinning models. The model FRÉCHETTE, M. 2000. Body size-density relationship for Mytilus

suggested by Guiñez & Castilla (2001) introduces the effective edulis in an experimental food-regulated situation. Oikos, 90: 28– 42.

density term that allows comparison of the self-thinning expo- BEGON, M., FIRBANK, L. & WALL, R. 1986. Is there a

nent between different populations or species with different self-thinning rule for animal populations? Oikos, 46: 122–124.

crowding strategies. The use of image analysis techniques to COELLI, T. 1996. A guide to FRONTIER Version 4.1: a computer

determine the effective density, as applied in this study of program for stochastic frontier production and cost function

M. galloprovincialis, could improve the results of the model estimation. Working paper 96/07, Centre for efﬁciency and

because the measured value of the occupied area is more realistic productivity analysis, University of New England, Armidale,

than the parallelepiped projection. The image analysis technique Australia.

used in the present study is easy to apply to other species, and its DAMUTH, J. 1981. Population density and body size in mammals.

application could improve estimation of occupied area. This Nature, 290: 699– 700.

variable is essential in obtaining realistic patterns of density- FRÉCHETTE, M., AITKEN, A.E. & PAGÉ, L. 1992.

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suspension feeder: consequences for self-thinning relationships.

arranged in complex matrices. Marine Ecology Progress Series, 83: 55–62.

FRÉCHETTE, M. & LEFAIVRE, D. 1990. Discriminating between

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self-thinning relationships. Marine Ecology Progress Series, 65: 15–23.

GUIÑEZ, R. & CASTILLA, J.C. 1999. A tridimensional self-thinning

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GUIÑEZ, R. & CASTILLA, J.C. 2001. An allometric tridimensional

model of self-thinning for a gregarious tunicate. Ecology, 82:

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GUIÑEZ, R., PETRAITIS, P.S. & CASTILLA, J.C. 2005. Layering,

the effective density of mussels and mass-density boundary curves.

Oikos, 110: 186 –190.

HUGHES, R.N. & GRIFFITHS, C.L. 1988. Self-thinning in

Figure 3. Representation of the distribution of mussels on the barnacles and mussels: the geometry of packing. American Naturalist,

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69: 531–534. ZHANG, L., BI, H., GOVE, J.H. & HEATH, L.S. 2005.

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territory size and metabolic rate as predictors of self-thinning in boundary line. Canadian Journal of Forest Research, 35: 1507–1514.

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