# Review and comparison of methods to study the contribution of variables in artiﬁcial neural network models

Convinced by the predictive quality of artificial neural network (ANN) models in ecology, we have turned our interests to their explanatory capacities. Seven methods which can give the relative contribution and/or the contribution profile of the input factors were compared: (i) the ‘PaD’ (for Partial Derivatives) method consists in a calculation of the partial derivatives of the output according to the input variables; (ii) the ‘Weights’ method is a computation using the connection weights; (iii) the ‘Perturb’ method corresponds to a perturbation of the input variables; (iv) the ‘Profile’ method is a successive variation of one input variable while the others are kept constant at a fixed value; (v) the ‘classical stepwise’ method is an observation of the change in the error value when an adding (forward) or an elimination (backward) step of the input variables is operated; (vi) ‘Improved stepwise a’ uses the same principle as the classical stepwise, but the elimination of the input occurs when the network is trained, the connection weights corresponding to the input variable studied is also eliminated; (vii) ‘Improved stepwise b’ involves the network being trained and fixed step by step, one input variable at its mean value to note the consequences on the error. The data tested in this study concerns the prediction of the density of brown trout spawning redds using habitat characteristics. The PaD method was found to be the most useful as it gave the most complete results, followed by the Profile method that gave the contribution profile of the input variables. The Perturb method allowed a good classification of the input parameters as well as the Weights method that has been simplified but these two methods lack stability. Next came the two improved stepwise methods (a and b) that both gave exactly the same result but the contributions were not sufficiently expressed. Finally, the classical stepwise methods gave the poorest results.

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