individuation follow from the fact that, like autocatalytic systems, ecological systems are more directly determined by external boundary conditions and material constraints than more complex, autonomous systems, such as organisms. Nonetheless, ecological systems can realize a basic regime of closure, just as autocatalytic sets.

An idea that is quite helpful when discussing the individuation of ecological systems is Montévil and Mossio's (2015) "tendency to closure." Closure offers a clear-cut criterion for drawing the boundary between a biological entity and its environment, providing a fitting solution to the problem of individuation, as the set of constraints subject to closure defines the system, based on the topological property of circularity in the network of constitutive interactions, whereas all other constraints acting on the system belong to its environment. Montévil and Mossio claim that we should ascribe closure to "maximally closed systems," i.e., systems including all mutually dependent constraints in the currently available descriptions (which are, by necessity, incomplete). Thus, in the case of mutually dependent organisms, there still seems to be a fundamental organizational continuity between the interacting organisms. In this case, it seems justifiable to ascribe ecological functions to the organisms constituting the system, even if the system does not show fully-fledged functional integration or constraints closure. Montévil and Mossio were discussing cases in which an encompassing system (say, a symbiotic one) is maximally closed, such that one might say that the symbionts within that system display a tendency to closure. As they depend on each other, they are not closed strictly speaking, but one can say they "tend" to be closed. We think we can extend this notion, however, to conceive of subsystems or modules, generally speaking, which show a relatively large degree of internal cohesion but yet depend on other modules in a given network. Closure ascription can extend in this case beyond each module, insofar as a maximally closed system should include all known constraints showing the topological circular property. Yet, we can claim that the modules containing – in the case of ecological systems – functionally coupled organisms or other items of biodiversity show a tendency to closure, as elements within a hierarchical set or network of modules. That is, in this case we can introduce a somewhat more relaxed notion of internal cohesion that makes it clear how the scope of the organizational theory of ecological functions is substantially broader than just a limited number of cases showing fully-fledged closure. To make this notion more precise, we can introduce a measure of the degree of closure in a system, based on the number of constraints that are both enabling and dependent, and, accordingly, are subject to closure. A tendency to closure points, then, to a specific degree of closure measured by the number of mutually dependent constraints in a system.

Ecological systems realizing closure of constraints can indeed exhibit different degrees of functional integration. They can be rather closed systems like the phyto-telmata of bromeliads, chosen by Nunes-Neto et al. (2014) as a case to develop the organizational theory of ecological functions, but not as exhausting all the possible cases to which the theory applies. They can be symbiotic systems or other functionally integrated consortia of organisms, in which control is exerted not only within but also across biological systems (Bich, 2019), as, for instance, bacterial biofilms
in which bacteria exchange enzymes (or DNA sequences coding for enzymes) responsible for the control of the internal metabolic processes in response to nutritional and other kinds of stress (Davey & O'Toole, 2000), or plants integrated by mycorrhizal networks that not only exchange metabolites but mutually affect their physiology and ecology (Selosse et al., 2006). These cases are different from the bromeliad one because a new order of functional integration is realized through the control exerted by organisms upon one another's processes.<fnref n="14" />

Ecological systems may show, however, much less integrated and bounded configurations and, yet, realize closure of constraints, as it happens when organisms exert control upon the conditions of existence of one another, either by directly harnessing the external flow of matter and energy or indirectly generating external control constraints in the environment, such as bird nests, spider webs, beaver dams, ant nests, etc. It seems clear, thus, that the domain of the organizational theory is much larger than some critics have supposed. There are plenty of systems in which ecological functions can be naturalistically grounded, in their teleology and normality, using the organizational theory.

### 13.4.3 Modularity Analysis and the Identification of Ecological Systems Showing Tendency to Closure

Surely, it is rather challenging to individuate ecological systems not as bounded as phytotelmata or beaver dams. Plant stands integrated by mycorrhizal networks, for instance, are difficult cases. However, we see this not as a fatal conceptual pitfall that the theory cannot deal with. It is rather a methodological challenge that can be tackled with its resources. Even though this is not the space to fully develop a methodological solution to the problem, we can advance some basic ideas on how to pursue it.

An analysis of modularity in ecological networks can provide at least an initial approach to identify ecological systems showing tendency to closure. Modularity – which describes the existence of subcommunities within networks – is currently regarded as a recurrent structure of many types of ecological networks (Thébault, 2013). A network shows modular structure when it consists of interconnected modules, while the extent to which species interactions are organized into modules amounts to the modularity of the network. In turn, a module in an ecological network is defined as a group of species more closely connected to each other than to species in other modules.

<footnote n="14">¹⁴Importantly, as Bich (2019) argues, the realization of a new order of functional integration does not imply that the organisms involved are not able to realize organizational closure and achieve functional integration by themselves. It just means that, while maintaining closure as functionally cohesive entities, they extend their functional networks of control constraints by realizing nested forms of functional integration that include more than one system and, we add, can also realize closure at a higher order.</footnote>
In an influential paper, Olesen et al. (2007) provide a good example of the relevance of modularity analysis for understanding the structure and functioning of ecological networks, given that modularity is both a key ingredient of network complexity and plays a critical role in their functioning, e.g., in relation to species coexistence and community stability. Indeed, the modular structure of species interactions in mutualistic networks was shown, for instance, to hinder species loss and promote long-term persistence of ecological communities (Krause et al., 2003; Kashtan & Alon, 2005; Olesen et al., 2007; Guimerà et al., 2010; Stouffer & Bascompte, 2011; Gilarranz et al., 2017; Sheykhali et al., 2020).

There are several underlying processes that can explain why ecological networks show modularity, all of which can be included in non-equilibrium models: modularity may reflect habitat heterogeneity, divergent selection regimes, and phylogenetic clustering of closely related species (Lewinsohn et al., 2006). It can also result from the convergence of species on correlated suites of traits shaped by similar interaction patterns, as captured by a concept commonly used in studies on plant-animal interactions, namely, that of syndromes (Fenster et al., 2004; Olesen et al., 2007; Dellinger, 2020).<fnref n="15" />

Modularity is no exceptional feature of ecological networks but rather a manifestation of a common property in biological networks, which, as Kashtan and Alon (2005, p. 13773) argue, “are modular with a design that can be separated into units that perform almost independently.” We can advance, thus, that modularity analysis can provide a first step to identify highly connected groups of species that may satisfy the requirements for showing tendency to closure.<fnref n="16" /> That is, organizationally closed (sub)systems<fnref n="17" /> in an ecological network can be searched for through the identification of modules, and the search space for those (sub)systems will be significantly reduced if we focus on modules of ecosystem parts that are more closely connected to one another than to parts included in other modules. After all, within a module, it is more likely that biological organisms/populations/functional groups will show mutual dependence due to their interactions, which are stronger than the interactions with other network components, i.e., it is more likely that they rely on one another for their own maintenance, with at least part of them being possibly both enabling and dependent constraints and, thus, being subject to closure.

The identification of modules in an ecological network can provide, thus, a first step to model organizationally closed ecological systems but needs to be complemented by an approach to investigate the within-module connections in order to

<footnote n="15">¹⁵ In fact, many pollination studies implicitly assume modularity when they focus on groups of interacting species sharing a syndrome.</footnote>

<footnote n="16">¹⁶ Although we cannot develop the argument in the confines of this chapter, we advance that the approach described in the body of the text may provide a way of implementing the procedure to delimit organizationally closed systems through the drawing of their spatial boundaries derived by Montévil and Mossio (2015) from the quantitative assessment of the tendency of constraints to be “packed together” in space.</footnote>

<footnote n="17">¹⁷ We write “(sub)systems” to accommodate the fact that the whole ecological network or more inclusive parts of it may be also described as systems in a number of cases.</footnote>
establish whether they take place between constraints and, moreover, between con-
straints that are both enabling and dependent. A possible way to model modules in
ecological networks as organizationally closed systems is to ascertain whether they
can be treated as ecological autocatalytic sets, as proposed by Cazzolla Gatti et al.
(2017, 2018). Another way, which we are currently investigating, is to show that
systems of differential equations used to describe coupled dynamics (e.g., consumer-
resources, predator-prey) can provide a mathematical framework to model ecosystem
closure of constraints. Surely, these two approaches can be integrated, as they
offer descriptions of the same dynamical system, with networks describing the
topology of the interactions, and differential equations, the dynamics of the
interactions.

Olesen et al. (2007) analyzed a total of 51 pollination networks, encompassing
almost 10,000 species of plants and flower-visiting animals and 20,000 links, and
found that 29 of them (57%) were significantly modular.<fnref n="18" /> In particular, all networks
containing more than 150 species were modular, while all those with less than 50
species were nonmodular. The modular networks had, on average, 8.8 ± 3.7 modules, ranging from a maximum of 19 to a minimum of 5 modules. Most links in such
networks were among species within the same module (on average 60% of all links),
reinforcing how modularity analysis may allow us to identify organizationally
closed (sub)systems in an ecological network, despite the intricacy of ecological
relationships and the relative openness of such systems. Individual modules in the
networks differed in size and shape because of both the variation in species number
and the ratio between pollinator and plant species. A module contained on average
32 ± 34 species (on average, 26 pollinator species and 6 plant species). This suggests that the set of organizationally closed modules or (sub)systems in ecological
systems may not be as small as some critics think. It was even the case that 36 (14%)
of all 254 modules identified in the networks were isolated species groups without
any links to the remaining network. However, this finding concerns the ecological
interactions between plants and animals modeled in the networks, and there is no
reason to assume that if other kinds of ecological relationships were at stake, those
same species groups would be equally isolated. Only 21 of these isolates, i.e., 4% of
all identified modules, were small 1:1 modules, consisting of only one pollinator
species interacting with one plant species. That is, just a minority of the modules
were the sort of small ecological systems, with just a couple of populations, that
Lean (2021) argued would exhaust most of the domain to which the organizational
theory could apply. Twenty-nine (11%) of all modules were star-shaped, consisting
of one generalist hub species, most often a plant species, showing no links to other

<footnote n="18">¹⁸ Olesen et al. (2007) treated all flower-visiting animals as pollinators, which, of course, is not necessarily true as several species may visit flowers without being involved in pollination but in other processes, such as nectar robbing. As the role of a species in an ecological network is defined by its topological position compared to other species, it is not central to functional ascription based on modularity analysis if the species at stake is a pollinator or not, since it may constrain the flow of energy and matter in a variety of ways and, thus, play different ecological functions according to the organizational theory.</footnote>
modules, while it was linked to a range of 3–51 peripheral pollinator species con-
nected only to the hub. Most of the hubs (189, i.e., 74%), however, varied a lot in
size and shape, showing the diversity of arrangements possible in plant-pollinator
networks. Some modules contained a set of species with convergent traits related to
their pollination biology, i.e., to pollination syndromes, or which were closely
related taxonomically.

Considering functional analysis, a rather interesting aspect of the study carried
out by Olesen et al. (2007) lies in the topological analysis of the role played by each
species in the networks. This role is defined by its position compared with other
species in its own module and how well it connects to species in other modules.
Accordingly, the analysis considers the relation between each species’ within-
module degree z, i.e., its standardized number of links to other species in the same
module and its among-module connectivity c, i.e., the level to which the species was
linked to other modules. Eighty-five percent of all species showed low z and c and
were peripheral species or specialists, showing only a few links and almost always
only to species within their module (72% of them had c = 0, with no links outside
their own module). Species with either a high z or a high c value were generalists
(15%), including module hubs (3%), i.e., highly connected species linked to many
species within their own module (high z, low c), and connectors linking several
modules (low z, high c) (11%). Species with high z and high c were network hubs
or super generalists (1%), acting as both connectors and module hubs. Plants were
the strongest module hubs. Connectors were mainly beetles, flies, and small-to-
medium-sized bees, and most network hub pollinators were social bees, especially
*Apis* spp. and *Bombus* spp., or large solitary bees, e.g., *Xylocopa* sp. and a few
*Diptera* species. Even though generalists not only contribute to pack peripheral spe-
cies together into modules but also connect modules together into networks, blur-
ring in this way module boundaries, it is possible to extract modules from networks
using the appropriate analytic approaches, as shown by several studies (e.g., Olesen
et al., 2007; Fortuna et al., 2010; Thébault & Fontaine, 2010; Schleuning et al.,
2014; Grilli et al., 2016; Sheykhali et al., 2020). This is instructive when one seeks
to consider how system openness does not entail that organizationally closed sys-
tems cannot be identified.

Some ecological networks may show a greater tendency toward modularity than
others, since this property is expected to increase with link specificity (Lewinsohn
et al., 2006).<fnref n="19" /> One may expect, for instance, that modularity is stronger in insect
herbivory or host-parasitoid networks, which show high link specificity, than in

<footnote n="19">¹⁹Link specificity concerns the degree of specificity of the ecological interaction represented by a certain edge in a network. For instance, in a food network, the more specialized the trophic relationship considered, the higher the link specificity, while the reverse is true for generalist trophic relationships. Link specificity is related to another key concept in the literature on ecological networks, namely, interaction intimacy (i.e., the degree of biological integration between interacting individuals; see Pires & Guimarães, 2013), such that the decision on specificity does not merely involve an analysis of the links in a given network.</footnote>
pollination and seed-dispersal networks, characterized by lower interaction speci-
ficity, and in traditional food webs (Olesen et al., 2007).

Important consequences for conservation may follow from the combined use of an organizational theory of ecological functions and modularity analysis, as it may allow us to ascertain key groups of taxa that need to be conserved for an ecological network to persist, based on the implications of losing them to the network functioning per se, not just on the choices of a scientist in relation to where to focus his or her attention. This is so because such a theoretical-methodological approach can provide us with normativity criteria that can underlie conservation decisions, for instance, about the conservation of biodiversity items that constrain the flow of matter and energy in the ecosystem in such a manner that its resilience and persistence – as aspects of its stability and, accordingly, of its intrinsic teleology – be maintained. These are criteria that depend on the natural normativity of the system and cannot be offered by accounts that fall short from grounding the teleology and normativity of functions. Consider, say, how the network consequences of species extinctions depend, among other factors, on the species role in the topology of the network. For instance, the extinction of a module hub may cause its module to fragment with no or minor cascading impacts on other modules, whereas if connectors are extinct, this may cause the entire network to fragment into isolated modules but with minor impacts on the internal structure of individual modules (Olesen et al., 2007). Accordingly, we can derive criteria, for instance, for choosing conservation priorities from the ascription of functions to different species depending on their topological roles in relation to the modularity of the network, which may be properly captured by interpreting ecological functions in terms of the organizational theory. To briefly mention a central topic discussed by Lean (2021), this will have normative consequences to decisions in invasion biology: for instance, alien invaders of a network may cause, as they are often highly generalist, fusion of modules in an ecological network, with profound, long-term effects on network functioning and selection regime (Olesen et al., 2007). This would be a reason, then, to choose to avoid the establishment of highly generalist invaders in ecological systems.

By considering the modularity of ecological networks, we can conclude, first,
that it may provide a first step in the identification of organizationally closed sys-
tems individuated as modules in a network (if complemented by approaches to
establish that the nodes in a module form a closure of constraints), which do not
necessarily correspond to a small set of small networks; second, that if we consider
the modularity of entire ecological networks, say, all species interacting through
ecological processes such as pollination in a given area, many networks will likely
include many modules, and, then, the fact that it may not be possible to describe the
whole network as a single organizationally closed system may not hamper func-
tional ascription based on the organizational theory within identified modules; and
third, that several different functional roles can be described based on modularity
analysis, such as module and network hubs, and that it may be possible to interpret
them based on the organizational theory, as related to the constraining actions of
biodiversity items on flows of matter and energy through, say, trophic or pollination
relationships.<fnref n="20" /> Even connectors, which link several modules, can be ascribed functions based on the organizational theory. In particular, this will be so if we consider coupling between modules as a functional role, something that will be possible if these connectors establish a form of mutual dependence between modules, due to their stable connecting interactions, such that one can say the self-maintenance of the modules is related to these interactions. Finally, the roles played by different species in the topology of ecological networks have conservation implications that are not at the discretion of scientists’ decisions only but also depend on the nature and structure of the networks, as modularity can even be said to spontaneously evolve in them (Kashtan & Alon, 2005).

## 13.5 Organizational Functions and Evolution

Dussault and Bouchard (2017) argue that the organizational theory dissociates the concept of ecological function from evolutionary considerations. It is true, on the one hand, that there is much work to be done in order to develop the connection between the causal loop by which functions explain the presence of the function bearer in an organization-based account, which has self-maintenance as its telos, and an evolution-based causal loop, which refers to the (past) natural selection of fitness-enhancing traits. But, on the other hand, the organizational theory has the resources to further develop this connection to the etiological dimension, which is part of its elaboration since its inception. The organizational theory proposed by Mossio and colleagues (2009; see also Moreno & Mossio, 2015; Saborido et al., 2011, 2016, among others) aims at accounting for the *explananda* of both etiological and dispositional theories of function. Moreover, in the theory of biological organisms under construction by the ORGANISM-group, which is also an important theoretical framework for our proposal, the concept of function is connected to the principle of organization, and this is in turn integrated with evolutionary thinking through the principle of variation (Montévil et al., 2016; Mossio et al., 2016). This means that in this framework evolutionary changes in organization along time, both qualitative and quantitative, are necessarily integrated into the understanding of biological phenomena, including ecological ones.

Ecological functions have been recently conceived as central for integrating evolutionary and ecological perspectives on ecosystems. As a consequence, even though the concept of function has played an important role in the whole history of ecology, it has become increasingly fundamental to the development of ecological and conservation research in the last three decades (Nunes-Neto et al., 2016a). This happened as a consequence of the biodiversity and ecosystem function (BEF) research program, which aims at establishing a better understanding of the

<footnote n="20">²⁰ In this sense, our arguments are not affected by the fact that not all flower-visiting animals in the pollination networks are truly pollinators but play different roles, for instance, as nectar-robbers, as observed above.</footnote>
relationships between biodiversity and the functioning of ecosystems (e.g., Naeem, 2002b; Loreau, 2010a, b). To account for these relationships, this research program attributes to functional diversity the role of a conceptual bridge between community and ecosystem ecology, i.e., between the understanding of biotic communities and, accordingly, of biodiversity, including the interactions among their constituting components and their effects, on the one hand, and their contribution to ecological processes that maintain ecosystems and their properties, on the other. This unification of ecosystem and community ecology is often recognized as an important goal both for the development of ecological knowledge and for meeting the challenges of the current socioenvironmental crisis (e.g., Pickett et al., 2007; Dussault & Bouchard, 2017). They are sought after by BEF researchers through the investigation of how specific traits of organisms and other biological items contribute to the maintenance and functioning of ecosystems. This requires, however, that the understanding of ecological functions be connected with how organismic functions are conceived in evolutionary theory, which is a central component of the theoretical framework in community ecology (Dussault & Bouchard, 2017). Not surprisingly, BEF researchers stressed that a synthesis of community and ecosystem ecology demands that evolutionary considerations be reintroduced into ecosystem studies (e.g., Loreau, 2010a, b).

It is at the purview of the organizational theory of ecological functions to deliver an understanding of functions that is both ahistorical and evolutionarily grounded. To do so, it will be necessary to elaborate more on the relations between the evolution of organisms and the emergence of ecological interactions and functions in organizationally closed ecological systems.<fnref n="21" /> A key aspect to bear in mind is that, as ecological systems emerge from interactions (at least part of them functional interactions) between populations that have been selected to a considerable extent, an integration between evolution- and organization-based accounts is a sine qua non for understanding ecological functions. But how should this integration take place? A fundamental requisite is to consider how to prioritize functional approaches in organisms or ecological systems. At the level of organisms, an evolution-based functional and teleological understanding should be grounded on an organization-based functional and teleological conception of self-maintaining organisms capable of survival and reproduction (Mossio & Bich, 2017). However, at the ecosystem level, it seems to be the case that organization-based function and teleology should be grounded on the interrelations among organismic functions (and also accidental or fortuitous effects) that emerge in evolution at the population level. That is, at the ecosystem level, organization arises from interactions among populations that have been selected for at the population level, as a kind of by-product of organismic functioning to achieve self-maintenance and increase fitness. After all, other populations are always a relevant part of the environment of any population at stake.

<footnote n="21">²¹ A dialogue with the persistence enhancing propensity (PEP) account proposed by Dussault and Bouchard (2017) can be helpful in this effort.</footnote>
Functions in ecology are relational and contextual, as emphasized by Dussault and Bouchard (2017). They emerge from current interactions between populations that are, at least partly, associated with organismic traits exhibiting functional roles that evolved historically, before a particular ecological system has been formed. This does not mean, however, that such functional traits have evolved for the sake of the ecosystem; rather, they partly evolved due to their fitness-enhancing consequences at the population level, partly due to other evolutionary processes than natural selection, and have been coopted for functional roles in ecological processes within the ecosystem when subject to its closed organization of constraints. For example, in plant-pollinator networks, different populations mutually stabilize each other (allowing for an account in terms of closure of constraints and organizational functions), but it is the evolution at the population level that explains the spread and eventual fixation of functions that are entangled with one another in the ecosystem closure of constraints (e.g., Patiny, 2012). Moreover, the historical constitution of ecosystems involves a “fine-tuning” of functional relations as a result of evolutionary paths, reinforcing the need to integrate evolutionary and organizational perspectives on ecological functions.

## 13.6 Ascribing Functions to Abiotic Items

For Dussault and Bouchard (2017), the PEP account accommodates the ascription of functions to abiotic components of ecosystems better than the organizational theory, as it allows function ascription to abiotic factors like disturbance regimes and habitat heterogeneity (e.g., Pickett et al., 1999; White et al., 1999). A similar argument is proposed by Odenbaugh (2019) but in a defense of a systemic approach, which is one version of a dispositional theory. But consider how we recently broadened the range of functional items that fall under the umbrella of the organizational theory in order to include abiotic items, provided they are under the control of bio-diversity items (El-Hani & Nunes-Neto, 2020). From this perspective, factors like disturbance regimes (say, related to fire) and habitat heterogeneity (for instance, due to the construction of a beaver dam) only play a functional role in an ecological system if they are products of constraints subject to closure in that system and are themselves involved in the production of constraints. If they are not so, then they are not truly functional but just boundary conditions that affect the maintenance of populations and ecological communities in that system (even though they can be functional if we are rather modeling an ecological system at a higher scale). If, say, habitat heterogeneity and disturbance regimes are not under the control of components of an ecological system, they do not have a functional role according to the organizational theory precisely because they are not under the control of the system and do not enable the conditions of existence of other constraints. But this does deny their relevance to the system's dynamics, as external boundary conditions. This relevance seems to be the reason why ecologists ascribe in a number of cases functions to such external entities and processes. A philosophical analysis can offer,
then, an important clarification about a conflation between the functional contribu-
tion from a system’s component and the dynamic relevance of external boundary
conditions (at a given scale). To describe the interaction of an external entity or
process not controlled by a system but influencing its dynamics in terms of a broad-
ened regime of functional integration is incorrect precisely because in this case the
system is not exerting any influence upon the generation of the boundary condition
(Bich, 2019). If we consider some ecological systems as showing the same kind of
regime of closure as autocatalytic sets (see, e.g., Cazzolla Gatti et al., 2017, 2018),
it will be also clear why one should not, from this perspective, ascribe an ecological
function to entities or processes external to the system and not under its control.
This conflates being a boundary condition to the system’s dynamics with playing a
function, which is an important drawback, since functions are attributed to compo-
nents of a given system.

If an external entity or process is under the control of the system, as disturbance regimes or habitat heterogeneity in a number of cases are, then it has a functional role defined in accordance with the closure of constraints defining the system, since by being under the control of the system, it becomes subject to closure, being both a dependent and an enabling constraint. Consider, as a case in point, how fire (as a disturbance regime), when integrated into the dynamics of an ecological system, say, through fire-adapted plant species exhibiting traits that promote flammability and, thus, influence fire frequency (e.g., Mutch, 1970; Schwilk & Ackerly, 2001), is not merely destructive but rather enabling, leading to regrowth processes that are crucial to the system's dynamics. In these cases, vegetation is a driver of fire regimes, and one can even talk about coevolution of fire and biota (McLauchlan et al., 2020).

Dussault and Bouchard (2017) consider, in fact, precisely the argument we are
advancing here but refuse it because it would, they argue, run counter to the ten-
dency in contemporary ecosystem ecology to include disturbance regimes into the
dynamic of ecosystems irrespective of whether they are under biotic control or not.
However, we think there is no real problem in this case, because boundary condi-
tions are part of the dynamics of the system no matter if they are within the closure
of constraints or not. This is a clear case in which the concept of closure of con-
straints is not properly expressed, since it only demands that part of the constraints
exerting influence on the system be internal to closure. A boundary condition that
affects the system without being within closure is still part of the system's dynamics.

Another argument presented by Dussault and Bouchard concern the difficulty of
determining whether a disturbance regime is under the control of the ecological
system, since it may lie on a continuum between being biotically controlled and
uncontrolled (Pickett & White, 1985, pp. 8–9). First, this is an empirical problem
that does not challenge the organizational theory: it is quite common in scientific
research that the stipulations of a theory pose empirical challenges for their applica-
tion to real-world processes. Second, the organizational theory can accommodate
through the idea of tendency to closure a situation in which an external process or
entity (say, fire) is somewhere on a continuum between being biotically controlled
or not. In this case we would in fact ascribe function to fire if it is at least to some
extent under biotic control.
Differently from Dussault and Bouchard (2017), who intend to follow ecologists' ascription of ecological function to abiotic entities and processes, no matter if they are under the control of the ecosystem or not, we rather think to be preferable to conceptually clarify the case from a philosophical perspective. It seems to us that – revisionist or not – the philosophical analysis at stake drives home a relevant distinction to ecological research, which we exemplify using fire as an example. If fire is under control of constraints internal to the ecological system, it can be both enabling and dependent, being part of the ecosystem closure of constraints, and, thus, being ascribed function, but if it is not under the control of those constraints, even if fire may be eventually enabling, it will not be dependent on internal constraints, and, thus, it will be just an external boundary condition, which should not be described as functional within that ecosystem, despite their significance to the system's dynamics. We do not see a problem in adopting a "revisionist stance" (as Dussault & Bouchard, 2017, p. 1133, calls it) in relation to some attributions of functions by ecologists. After all, epistemological studies would be quite limited in their utility and contribution to scientific research if we assumed that philosophical analysis could never clarify the uses of concepts by scientists themselves.

It does not matter, then, if some abiotic process has the same effect on an ecosystem as a biotic process to which an ecological function is ascribed, as in the example of nitrogen fixation by lightning or volcanoes. Contra Odenbaugh (2010, p. 251), this does not mean that those abiotic processes should be ascribed a function as well, since what they have in common with the biotic process at stake is just that they are both boundary conditions. Nevertheless, the crucial distinction between being a boundary condition under control or deprived of control by the system still applies and is, in our view, crucial to keep in place the distinction between what is truly functional and what merely affects the system's dynamics. From the perspective of the ecological system, nitrogen fixation by a lightning is merely a boundary condition (which, by accident, can fortuitously affect an ecosystem's dynamics, or eventually become stable enough to affect the dynamics on a steady basis<fnref n="22" />), while the same process carried out by bacteria has an ecological function.

Another case that does not bring as much trouble to the theory as Dussault and Bouchard (2017) think concerns source-sink dynamics (e.g., Pulliam, 1988; Pulliam & Danielson, 1991; Amarasekare & Nisbet, 2001; Loreau et al., 2003), which explains the maintenance of “sink” populations, i.e., populations which would run locally extinct if not maintained by constant immigration from “source” populations, as well as of “source” populations whose abundance would often inflate considerably if there was no emigration to “sink” populations. They correctly argue that the effects of source-sink dynamics are often indistinguishable from those of more conventional density-dependent regulation processes (as discussed by Sterelny, 2006, pp. 219–220) and would thus warrant ecological functional ascriptions just as

<footnote n="22">²² In this case, it may be that the dynamics of the system eventually turns the boundary condition subject to closure, meaning that the abiotic process become a functional part of that system. This does not affect, however, the core of our argument.</footnote>
in the latter case. True. But there seems to be no problem, however, in ascribing
ecological functions to sink and source populations based on the organizational
theory. It is only necessary to describe a higher-level entity of which those popula-
tions are part, playing functions within its closure, such as a metacommunity or
metapopulation, depending on whether we are dealing with multiple or single spe-
cies. Therefore, in the case of both density-dependent regulation processes and sink-
source dynamics, the closure criterion can be met, and function can be ascribed
according to the organizational theory.

13.7 A Word on Pluralism About Ecological Functions

It is worth saying here a few words on pluralism about functions. In the overall literature on biological functions, pluralism has been often regarded as an attractive option. For instance, a number of authors supported a pluralistic solution to the problem of function by advocating that the etiological and dispositional theories offered two complementary concepts (e.g., Millikan, 1989; Amundson & Lauder, 1994; Allen & Bekoff, 1995). Godfrey-Smith (1993) called this solution a “consensus without unity.” Currently, one cannot advocate for pluralism about functions without considering also organizational theories, in their several versions, as one of the key players in the debate. Specifically in the ecological domain, Garson (2018) has also defended within-discipline pluralism about functions.

But, when we are dealing with some specific problem, it seems to us that plural-
ism should be the conclusion we reach once we did our best to find a single, unify-
ing theory. It should be the outcome of an investigation that justifies the principled
impossibility of a unified account. If we think that a certain theory about function,
ecological or otherwise, cannot be the unique one, a proper justification should be
offered. Why is it the case? Moreover, to avoid empty pluralism, we need to identify
which kind of phenomena can be accounted for by which models, such that we may
in the end reach a theory unified as a family of models, as proposed in a pragmatic
view of theories in ecology (Travassos-Britto et al., 2021).

To our understanding, we are not yet at a point in the investigation that allows us
to settle the case and conclude for a pluralistic perspective on theories of ecological
function. The jury is still out. Thus, rather than assuming pluralism, we will leave
for now this possibility open and continue inquiring into the application of the orga-
nizational theory to different uses of functional explanations and ascriptions in eco-
logical and conservation research. This does not mean that we are claiming that
normative functions will be properly attributed to each and every ecological system.
Also, this is not the same as exclusively defending organizational functions as an
overarching account for all functional ascriptions in ecological research, as Lean
(2021) claims to be our intention. We are simply continuing to pursue our avenue of
inquiry, extending the theory as much as we can, but open to the possibility that it
may not apply, eventually, to a number of functional ascriptions made by ecologists.
The organizational theory of ecological functions remains expansive to new cases, and we really do not know if the latter may be the case.

As an example of how the domain to which the organizational theory is applied
can be extended, we can consider two mechanisms proposed to explain how biodi-
versity enhances the maintenance and resilience of ecosystems, namely, sampling
and compensation effects (e.g., Sterelny, 2005).<fnref n="23" /> In the former mechanism, the
increased resilience of species-rich ecosystems is attributed to the statistical fact
that they have more chances to contain species whose functional performance will
not be affected by a range of environmental variations. In this case, the functional
contribution to ecosystem maintenance is attributed to items of biodiversity rather
than to biodiversity as a whole, and no difficulty is posed for the organizational
theory. Compensation effect is, however, a different matter, as the increased resil-
ience of species-rich ecosystems is related in this case to response diversity, i.e., the
presence of many species that respond differently to environmental variation but are
able to perform similar functional roles in the ecosystem. Response diversity entails,
thus, that the species may show compensatory dynamics, i.e., when an ecosystem is
subject to variation in its interaction with other systems that leads a formerly domi-
nant species to decrease in abundance, the functional consequence for the ecosys-
tem dynamics can be buffered by the compensation of another species that is
functionally equivalent but shows a differential response to the variation at stake.
Thus, the likelihood that a variation leads to impacts that may disrupt ecosystem
functioning and harm its capacity of maintaining itself is reduced, and, conversely,
ecosystem resilience before that variation is maintained or even enhanced.

Compensation effect illustrates a case in which more work is needed to extend
the domain of the organizational theory. In the definition of organizational function
proposed by Saborido et al. (2011), function is ascribed to a *trait* that exerts a con-
straint subject to closure in an organization of a given system. Mossio et al. (2013)
characterize a constraint as a *configuration* and Moreno and Mossio (2015) as an
*entity* that exhibit a symmetry with respect to a process (or set of processes) under
its influence. Nunes-Neto et al. (2014) consider *items of biodiversity* as objects of
functional ascription in ecology, while El-Hani and Nunes-Neto (2020) recently
broadened the set of functional objects in the theory to include *abiotic items*.
Biodiversity is a global property or, to put it differently, a distributed feature of an
ecological system. The question that arises is as follows: Can a global property be a
constraint, such that the organizational theory justifies the ascription of ecological
function to biodiversity *per se*? At this point, we do not see any fundamental block-
age for formulating the notion of constraint in such a manner that this justification
can be done. While it may stretch the concepts too far if we treat biodiversity as an
entity in an ecosystem, the use of categories such as trait and configuration in the
organizational functional discourse paves the way to encompass biodiversity as an
object of functional ascription. This will need some reworking of the notion of

<footnote n="23"><sup>23</sup>The arguments in this paragraph benefited a lot from the discussion made by Dussault and Bouchard (2017, pp. 1133–1134).</footnote>
constraint in order to include under its extension global properties such as biodiver-
sity, but this will not be some far-fetched conceptual operation. Therefore, while
there is still work to do, the organizational theory can be applied to explain both the
specific functional contributions of many items of biodiversity to the overall func-
tioning of an ecosystem and the collective stabilizing function of response
biodiversity.

13.8 From Organizational Functions to an Integrated
Scientific and Ethical Approach to Sustainability

The fortunes of teleological accounts of ecological functions, such as the organiza-
tional theory, may have important consequences to conservation ethics. If we show
that ecological systems are structured in such a manner that their parts are func-
tional for the whole, we may be able to provide support to the claim that they pos-
sess a type of natural value on a naturalistic basis. Such a natural normativity can
facilitate objective judgments about the role of populations within ecosystems and
about conservation measures, as well as mediate debates in conservation ethics and
provide guidance for thorny environmental ethical questions.

What does the idea that ecological (and, possibly, socioecological<fnref n="24" />) systems
realize closure of constraints entail, then, for an ethical perspective on such sys-
tems? Recognizing that biological systems include constraints that perform func-
tions is to recognize a normative dimension of the very existence of these systems.
In this sense we can differentiate between two kinds of systemic state, namely,
between organized states, which exist according to the norms of the system's behav-
ior, maintaining conditions of existence that allow its persistence and resilience, and
states that work counter to the system's organization, deviating from the norms of
its behavior and disrupting its conditions of existence (Moreno & Mossio, 2015;
Cooper et al., 2016; Montévil, 2021). That is, the functionality of certain biological
features concerns not only a current performance of an ecological system but a per-
formance that the system must do in order to continue to exist. It seems, then, that
we may be able to discern in a normative way between *good* and *bad* functioning of
ecological systems (Cooper et al., 2016). It is at this point that the descriptive lan-
guage of biological organization touches on ethical and axiological aspects.

Notice, however, that this is a more demanding normative dimension than that at
play when we just speak of functions. If we consider, say, a pumping heart, this
organ will be fulfilling the biological norms involved in the performance of its func-
tion even when pumping poorly, with consequences to the organism's health. To
consider the performance of the heart's function in a healthy condition demands,
thus, a second, additional set of norms, establishing that the heart is not only

<footnote n="24">²⁴For a brief and initial discussion of the application of the organizational theory of functions to socioecological systems, see Nunes-Neto et al. (2016b).</footnote>
functioning but also functioning *well*. Accordingly, one thing is ascribing normativ-
ity to ecological functions based on the intrinsic purposiveness associated with the
realization of closure by an ecosystem, as a causal regime maintaining its own con-
ditions of existence. Another thing is considering whether an ecosystem is function-
ing poorly or well, as this requires a second source of normativity. What should this
source of normativity be is one of the issues to be tackled by an environmental eth-
ics theory.

There are important differences, however, between proposing an organizational
view of ecological systems and functions (which is mostly an epistemological
stance) and developing an ethical perspective on them (which entails an interpreta-
tion based on moral philosophy). Let us begin, thus, by appreciating an important
conceptual difference in moral philosophy which is important for our arguments,
namely, that between moral agents and moral patients (Warnock, 1971; Goodpaster,
1978; Nunes-Neto & Conrado, 2021). A moral agent is a being capable of emitting
moral judgments, which can be – as a consequence – held responsible for its actions.
In turn, a moral patient is a being that matters in relation to actions and, accordingly,
should be taken into consideration in moral judgments about the latter. When deal-
ing with ecological systems, we are primarily talking about moral patients, rather
than moral agents, who could have any kind of moral duty, obligation, or responsi-
bility. When we refer to what an organism or species (say, a bee species) should do
in relation to the norms of an ecological system’s behavior (say, in a pollination
network), we are surely not considering any moral duty, obligation, or responsibility
but just manifesting an expectation that a given behavior must happen if those norms
are to be observed and conditions needed for the system’s resilience and persistence
are to be fulfilled. But this expectation may also offer criteria to distinguish between
what is good or poor working of the system, providing an ethical perspective on its
organization and functioning.

But would we not be committing a fallacy – namely, the naturalistic fallacy – by
constructing an ethical perspective on ecological systems from an organizational
theory of functions? We cannot simply make inferences from purely factual claims
to moral ones, or, to put it differently, *normative claims* about what *ought to be true*
can never be validly inferred from *factual claims* about *what is true* (e.g., Kitcher,
1993; Sterelny & Griffiths, 1999). This means that the use of good, bad, well,
poorly, or other normative terms in an ethical context does not entail merely an
expectation about the natural behavior of systems but also about what we consider
that we – human beings as moral agents – *must do* in relation to others (humans or
nonhumans), to moral patients, in our everyday life. There is a central difference
between developing an organizational theory of ecological functions and an ethical
perspective on ecological systems: while the normative language in the organiza-
tional theory expresses facts (even if dispositional), the normative language in the
ethical field expresses values. The kind of normativity that stems from the organiza-
tional theory does not come from the same sources than those at play with ethical
and moral human judgments. When we consider an ethical view about ecological
systems, we must also recognize, thus, our own (human, thus ethical) perspective,
which – at least from our point of view – cannot be reduced to a naturalized
outlook about organizational functions only. In this sense the difference between
ethics and natural sciences is of central relevance, even if a dichotomous view of
facts and values is avoided:<fnref n="25" /> ethical issues concern, preponderantly, matters of
value, while the natural sciences deal, mainly, with matters of fact, but matters of
value and fact, albeit not entailing one another, do interact.

This difference does not mean, however, that it is not possible to build an inte-
grated perspective combining the organizational theory and an ethical theory. It only
means that we should not do so by committing a naturalistic fallacy, since one thing
is an epistemological (naturalistic) outlook on organisms and ecosystems, and
another thing is an ethical standpoint. But these stances are not necessarily in con-
tradiction; they can interact with one another, and, perhaps, in some cases be even
conceived in a kind of continuity or complementarity. As Sterelny and Griffiths
(1999) argue, even if moral principles cannot be inferred from purely factual bio-
logical premises, we can discover morally relevant facts through biological research,
which can interact with existing moral principles to produce new practical policies.

In what follows, we are not going to talk about ethics in general, but rather talk
about environmental ethics, since ecological systems are our main focus here. As
soon as environmental problems gained notoriety (around the 1970s), a new field of
ethical reflections was consolidated, environmental ethics, as a way of dealing with
a whole range of new issues that could not be well grasped by more traditional ethi-
cal frameworks. This was so because those issues concerned a series of beings and
processes that had not been commonly considered in previous ethical studies. In its
emergence, environmental ethics differed from previous views, which were gener-
ally anthropocentric (i.e., focused on human beings). In this sense, environmental
ethics broadened the scope of ethical study and reflection to include other natural
entities and processes such as animals, plants, rivers, mountains, ecosystems, etc.
This was an expansion of the scope of moral considerability (i.e., concerning which
beings or entities should be morally considered in our decisions and actions). What
was at stake, in short, was which among all the natural beings should we humans (as
moral agents) accept as moral patients (Warnock, 1971; Goodpaster, 1978; Vaz &
Delfino, 2010).

Kant [1785] 2007) differentiated between direct and indirect moral consider-
ations, depending on the moral status we recognize in other beings. We consider
something to be under the purview of indirect moral consideration when its value is
not final but rather justified by reference to something else, which is external to it.
For example, the value of a hammer comes from the act of hammering, which is
external to the hammer itself. In this case, the value of a hammer is merely instru-
mental. In turn, we generally accept – in accordance with Kant’s view – that the
importance of a human life is final, in the sense that it has value in itself, without

<footnote n="25">²⁵Following Putnam (2002), we do not endorse a dichotomy between facts and values (as assumed, for instance, by logical positivists), but this does not mean that we cannot differentiate between them. Every fact is value-laden, as well as values are connected to facts in the empirical domain. Here, we assume a non-dichotomous difference between facts and values, as well as between science and ethics, recognizing at the same time that there are mutual influences between them.</footnote>
requiring justification in terms of anything else. Accordingly, a human life cannot be
grouped generically together with other entities that might supposedly replace it in
fulfilling some external value (as is the case of a hammer, which, when broken, can
be replaced by another one, with no harm to the satisfaction of its value). This
means a human life is irreplaceable and shows intrinsic value, i.e., a value that is
justified in itself. Direct moral consideration results from the recognition of this
type of value (Warnock, 1971; Goodpaster, 1978; Vaz & Delfino, 2010).

For Kant ([1785] 2007), only human beings – as rational beings – should have
their intrinsic value recognized, being fundamentally different, in ethical terms,
from things and other beings. However, the appraisal of this humanist position has
changed with the emergence of environmentalism, among other developments. This
view came to be regarded as a form of anthropocentrism. Environmental ethics
translated moral perceptions that came to the fore with environmentalism into the
proposal of expanding moral theories in such a manner that recognition of intrinsic
value in other beings could be justified (Warnock, 1971; Goodpaster, 1978).<fnref n="26" />

This expansion of moral theories resulted in a variety of different positions. The
sentiocentric current,<fnref n="27" /> for instance, attributes intrinsic value to all sentient beings,
i.e., to all those that can experience their own life (including humans and a range of
nonhuman animals). The biocentric current, in turn, recognizes the intrinsic value of
all living beings, whether they are sentient or not (also including, say, bacteria,
fungi, etc.). The ecocentric current, finally, ascribes intrinsic value to ecosystems
and cannot be regarded as a mere expansion of other moral theories, due to its more
holistic character (Vaz & Delfino, 2010; Nunes-Neto & Conrado, 2021).

In the wake of this theoretical expansion, one of the main tasks has been to jus-
tify the intrinsic value of nonhuman beings. This means to offer reasons to justify
which of these beings (if any) have a purpose of their own. As the organizational
theory discussed here naturalizes the concept of function in living systems, it offers
a possible contribution to the understanding of this purpose (see, e.g., Holm, 2017;
Moosavi, 2019). Biocentrism offers a case in point. In the case of this stance, the
justification for ascribing intrinsic value to all living beings stems from the idea that
the intrinsic teleology associated with organisms provides a criterion for objective
recognition of a good of its own, a good that does not originate from subjective
attribution of value (e.g., Taylor, 1986; Varner, 1998).

Holm (2017) investigates whether the biocentric claim can be well justified by
the organizational theory in response to what he calls the scope problem. According
to this problem, for the biocentric justification to correspond with the moral

<footnote n="26">²⁶We chose here, for simplicity, a Kantian way of describing the changes brought about by the emergence of environmental ethics. However, there are other equally important moral theories, such as utilitarian and virtue ethics theories, that would describe the research tradition of environmental ethics differently.</footnote>

<footnote n="27">²⁷Sentiocentric ethics can be understood not as an environmental ethics *per se* but as an animal ethics, with its own research agenda. However, it is part of the same movement of questioning and overcoming the anthropocentric position and that is why it is described here within the same tradition of environmental ethics.</footnote>
intuitions of biocentrists, it is necessary that the teleology identified in living beings encompasses all types of possible organisms and be exclusive to them, i.e., not shared with non-organisms (e.g., artifacts and inanimate objects in general). Holm argues that the organizational theory, to a large extent, locates the scope of teleology in the domain of living systems, except for the theoretical possibility that some dissipative systems, such as candle flames and hurricanes, also show a rudimentary sort of constraint closure, resembling the intrinsic teleology described by the organizational theory. He considers, then, that this possibility poses a problem for the defense of a strict view of biocentrism, as it more appropriately points to a defense of a teleocentrism, which acknowledges that beings that are not organisms can also (albeit arguably) be targets of direct moral consideration whenever they show intrinsic teleology.

Moreover, the naturalization of the ascription of functions to biological items, as articulated by the organizational theory, is regarded by Holm as suggesting a potential empirically testable criterion for the biocentric claim. That is, as any system realizing self-determination by means of a closure of constraints will exhibit intrinsic teleology and, hence, a good of its own, the organizational theory enables biocentrists to turn the claim that living systems show such a good into an empirical thesis, without appealing to the contested concept of “life.”

Holm's proposal of a teleocentrism points to the possibility that a supraorganis-
mic system be regarded as having a good of its own, provided it shares the same
kind of orientation toward the end of self-maintenance exhibited by organisms
(which awakens the moral feelings of biocentrists). Once we consider that this is the
case of ecosystems, the path is open for an ecocentric argument, such as that devel-
oped by Rolston, III (1987), who understands nature as a set of teleologies, ranging
from human self-legislation to ecosystem self-maintenance, passing through organic
autonomy.

Nunes-Neto et al. (2014) support this understanding by showing how an ecosys-
tem can be treated as an organizationally closed system in which the items of biodi-
versity (and abiotic items, see El-Hani & Nunes-Neto, 2020), acting as mutually
dependent constraints on the flow of matter and energy, give rise to intrinsic teleol-
ogy, just as we observe in organisms, even though ecosystems typically lack other
distinctive features of the latter, such as agency. Once one accepts the organizational
theory of ecological functions, it might seem that a strictly biocentric position could
not hold, since the same criteria for the good of organisms may be also valid for
ecosystems. However, it is not really the case, to our understanding, that the teleo-
logical grounding of ecocentrism on the organizational theory of ecological func-
tions denies the epistemological legitimacy of biocentrism. Looking more closely,
we must notice that, as Nunes-Neto and Conrado (2021) argue, biocentrism and
ecocentrism are not contiguous perspectives on the scope of moral considerability
but, instead, are focused on different kinds of entities. While biocentrism lies in the
same spectrum as, for instance, sentiocentrism and anthropocentrism, ecocentrism
is a response to the lack of moral considerability of nature in general, arguing that
holistic entities such as ecological systems should be morally considered as having
intrinsic value. This means that biocentric and ecocentric perspectives are not mutu-
ally exclusive, even though in some situations there could be tensions between
them, such as in the classical example of hunting wild animals for maintaining
ecological attributes of ecosystems (see, e.g., the debate between Regan, 2013, and
Callicott, 2010) or the example of cutting and removing a tree in order to produce
organic matter to maintain an agroforestry system (Miccolis et al., 2019).<fnref n="28" />

Following the argument above, if an ecological (or for that matter, socioecologi-
cal) system realizes closure of constraints in a similar way to organisms, then it will
be also a candidate for the recognition of its own good. What does that mean?
Namely, that each and every ecological (and socioecological) system would have its
own good, considering only the criteria provided by the realization of closure of
constraints and intrinsic teleology. However, this conclusion would lead to serious
moral conflicts since, if we dissociate the whole from the parts, it will be possible to
conceive the well-being of the whole, even if there is no well-being of one or more
parts. For example, it would be possible to think that a socioecological system
including slavery might have a good of its own if it showed organizational closure.
However, just as ecosystems are formed by items of biodiversity that exhibit their
own individual good, so are socioecological systems and, accordingly, the claim
that a socioecological system including slavery might have a good of its own would
not hold. Rather, we would be facing in this case a conflictive state of affairs. This
is analogous, in fact, to a dilemma discussed above: just as there may be conflicts
between biocentrism or sentiocentrism, on the one hand, and ecocentrism, on the
other, the same is true in the case of socioecological systems. These conflicts will
happen whenever there are tensions between the intrinsic goods of individual

<footnote n="28">²⁸ Another example of tension between biocentric and ecocentric perspectives concerns the impli-
cations of redundancy to conservation decisions based on considerations about role functions. For
instance, if two species play the same role function in an ecosystem and the extinction of one of
them does not impact sustainability (because of redundancy), functional considerations may fail to
provide a rationale to preserve it. This is a relevant problem for conservation decisions justified on
functional grounds, which does not go away when we propose, from an integrated scientific and
ethical point of view, a conception of sustainability that entails our duty as moral agents to support
the self-maintenance of ecological systems (see below). This is not the space to engage with this
issue in the depth it deserves, but let us just briefly state that, first, pluralism about functions may
play an interesting role in this respect, since conservation decisions that seem attractive, but are not
justified by some theory of ecological function, may well be justified by another one. Second, that
the problems entailed by redundancy for conservation decisions have been recognized and debated
in the scientific literature for a while, and one of the outcomes of the discussion has been the
requirements of more fine-grained descriptions of ecological role functions, such that what at first
may seem to be a redundancy may eventually be shown to be a case of functional complementarity
between the roles played by two or more species in relation to ecosystem processes (e.g., Rosenfeld,
2002; Oliver et al., 2015). An ecological community can only be maintained if there is functional
complementarity among several anatomic, physiological, behavioral, and other attributes of the
populations composing it, as it has been shown, for instance, in several studies on pollination sys-
tems (e.g., Brittain et al., 2013; Fründ et al., 2013). Surely, in the case of complementarity, the
conservation of all species at stake will be justified.</footnote>
organisms (humans or not) and the intrinsic goods of whole ecological or socioecological systems. It seems to us that these dilemmas can be avoided by an understanding of the system's well-being as integrated to the well-being of its parts, which seems reasonable, once the system is composed by the parts and their interactions. Cases of conflict between the system itself and its parts may generally involve some kind of malfunctional behavior. However, a more complete evaluation of this problem is out of the scope of this chapter, and we shall leave it for future investigation.

What does the recognition or ascription of intrinsic value to an ecological (or socioecological) system mean? First, that we consider that system as important in itself, that is, as having a purpose of its own, or a value of its own. Second, that we judge we have a duty to the system with regard to its self-maintenance. In short, we must promote the resilience and persistence of the system, and not its destruction. This is equivalent to saying that we must sustain the system, i.e., that we must act toward its being a sustainable system. In short, sustainability, from this perspective, is the realization of the duty to promote the good of an ecological (or socioecological) system that has its intrinsic value duly recognized by virtuous moral agents integrated into a worldview of respect for nature. This new conception of sustainability provides an alternative to the usual anthropocentric and economically based version, associated with the management of natural resources, (social, economic, or ecological) capital, and/or ecosystem services. By combining intrinsic valuation with the self-maintenance of ecological (or socioecological) systems, this new conception allows us to use a common “grammar” to refer to respect for nature and responsibility (see Larrère, 2013) in such a manner that the values of technological progress, capital, and the market can be subordinated to what Hugh Lacey (2014, 2016) calls “viable values,” associated with the sustainability of socioecological systems, social justice and participation, and universal well-being.

The organizational theory of ecological functions offers a promising way not only to further develop important positions in environmental ethics but also to integrate fields of ethical knowledge hitherto pursued in a relatively independent manner. This does not mean – it is important to notice – that the organizational theory can ground by itself an ethics. This theory, applied to organisms or ecological systems, offers a naturalized epistemological perspective on their organization and intrinsic teleology, which is not sufficient, in our view, to ground ethical aspects related to the interactions between human beings (as moral agents) and other beings or systems (as moral patients). These ethical aspects demand a consideration, both in theory and practice, of properly ethical and moral perspectives (for instance, theories providing criteria to ground the value of moral actions, or differences in value ascription), which cannot be reduced to a naturalized approach. Nonetheless, there is much to gain from an interaction between morally relevant features of a naturalized approach to the intrinsic teleology of organismic and supraorganismic systems and principles provided by ethical and moral theories.
## 13.9 Concluding Remarks

We further developed in this chapter the organizational theory of ecological functions by responding to some criticisms that allowed us to sharpen the theory. We argued about the individuation of ecosystems as organizationally closed systems, to which the theory can be applied, provided some comments on how evolutionary considerations may be integrated into an organizational understanding of ecological functions, and took additional steps for elaborating on how functions can be ascribed to abiotic items according to the theory. We expect to have shown how the organizational theory provides a convincing basis for naturalizing the teleological and normative dimensions of ecological functions, as well as for making contributions to the construction of an integrated scientific and ethical approach to sustainability that can avoid an anthropocentric perspective.

**Acknowledgments** We are indebted to Alan Love, Matteo Mossio, Gaëlle Pontarotti, and an anonymous reviewer for their comments on a previous version of the manuscript, which importantly contributed to its improvement. The work on this chapter has been supported by funding from the John Templeton Foundation (#62220) within the cohort program "Agency, Directionality & Function," coordinated by Alan Love (University of Minnesota, USA). We are also indebted to CNPq (proc. nº 465767/2014-1), CAPES (proc. nº 23038.000776/2017-54) and FAPESB (proc. nº INC0006/2019) for their support to the National Institute of Science and Technology in Interdisciplinary and Transdisciplinary Studies in Ecology and Evolution (INCT IN-TREE), in which the present study has been developed. NNN thanks CNPq for research funding (proc. nº 423767/2018-6). CNEH thanks CNPq for the concession of productivity in research grants (proc. nº 303011/2017-3 and 307223/2021-3) and CAPES/UFBA for senior visiting research grant (proc. nº 88887.465540/2019-00). FRGL is thankful to CAPES for doctoral studies grant (proc. nº 88887633926/2001-00).

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<figure><img src="image_42.png" /></figure>
# Index

## A

Adaptationism, 130, 140, 166,
221, 230

Agency, 11, 15, 16, 48, 77, 82,
87–98, 159–182, 201, 203,
219, 264, 266–268, 299, 319

Anthropocentrism, 285, 287, 295,
317–319, 321

Anticipation, 89–98

Aquinas, T., 25

Aristotle, 24–26, 32, 36, 42, 44, 67, 142,
178, 287

Autonomy, 29, 32, 36, 43, 47, 52,
89–99, 105–108, 110, 115–122,
164, 176, 178, 196, 198, 226,
232, 250, 253, 254,
265–267, 319

and robotics, 91

theory of, 10, 15

Autopoiesis, 13, 14, 32, 43, 133,
163, 193, 196, 208

## B

Baer, K.E. von, 12, 129

Bernard, C., 12, 28, 43, 49–51, 226

Bertalanffy, L. von, 2, 13, 33, 47,
106, 226

Biodiversity and ecosystem function (BEF)
research program, 294, 308

Biological specificity, 267

Boerhaave, H., 30

## C

Canguilhem, G., 44, 49, 50,
52–54, 60, 62, 67, 74,
78, 81, 266

Cartesianism, 69, 77–79

Category mistake, 44, 45, 47

Causal vs. non-causal determination, 109

Causal thicket, 161, 166–169

Cell

filopodia, 273

motility, 268, 274, 276, 277, 280

proliferation, 173, 268, 271,
275–277, 280

pseudopodia, 273

Central dogma of molecular biology, 2

χ symbol, 273, 274

Closure, 13, 14, 30, 31, 34–37, 268, 289,
300, 301

of constraints, 14, 107, 163–165,
174, 179, 181, 227, 228, 243,
250, 259, 268, 269, 289, 291–293,
296, 297, 299–303, 305, 307,
310, 311, 315, 319, 320

ecosystems, 254, 291–294, 305,
316, 319

organizational, 162–165, 174, 175,
179–181, 227, 243, 250, 251, 253,
259, 293, 294, 299, 300, 309,
310, 314, 320

Cognition, 85, 89–98

Collagen, 269, 271, 274, 277, 279, 280

fibers, 272–275, 277, 280
Complexity, 11
  functional, 8, 10, 11, 45, 206, 207, 251, 294, 301, 320
  organized, 9, 10

Constraint, 35, 36, 163-165, 192, 201, 205, 251, 267, 289, 291, 300, 305, 314
  functional, 10, 35, 53, 164, 165, 175, 203-205, 210, 226, 227, 229, 230, 250-252, 268, 276, 293, 297-299, 310, 312
  material, 35, 204, 207, 208, 250
  primary, 278
  regulatory, 208, 210, 251, 252, 259, 279
  trans-generational, 203, 204, 208, 210, 227

Cuvier, G., 12, 31, 130

Cybernetics, 7, 34, 177

## D

Darwin, C., 2, 31, 129, 159, 160, 165, 166, 175

Descartes, R., 26

Developmental resource, 168, 175

Developmental systems theory, 42, 128, 168, 224

Development/ontogenesis, 167, 171, 172
  constructive, 4, 5, 160
  principle of, 172

Dialectics, 135, 160, 163, 164, 171, 177, 179

Diderot, D., 28

Downward causation, 112, 118, 119

Dynamical
decoupling, 205-210, 212
presupposition, 164, 171, 175
systems theory, 93, 180

## E

Ecological networks, 303, 307

Ecology
  equilibrium, 295, 297
  non-equilibrium, 295-297, 304

Eigen's paradox, 169

Emergence, 4, 8, 9, 28, 87, 99, 164, 165, 181, 189, 192, 203, 204, 208, 274
  transformational, 114, 115, 117-122

Emergentism, 105, 111-113, 115, 119-122, 176, 177

Epigenesis, 80

Epigenetics, 134, 138-140, 142, 149, 168, 221, 224, 227, 231, 232

Epistemology, 28, 43, 44, 46-48, 51, 53, 54, 61, 63, 68, 72, 108, 133, 161, 177, 266, 267

Epithelium, 269, 270
  acinus, 273, 277
  duct, 273, 277

Error catastrophe, 170, 171

Ethics
  conservation, 295, 315
  environmental, 287, 315-318, 321

Evolution, 33-35
  Darwinian, 159, 165-166, 172, 181, 188
  and the struggle for existence, 159, 160, 166, 175, 181
  theory of, 2, 172, 175-177

Evolutionary developmental biology
  (evo-devo), 135, 160, 178, 180

Evolutionary synthesis, 161
  extended, 4, 5, 160, 177, 220-222
  modern, 1, 4, 159

Evolvability, 169-173, 221

Explanation
  lineage, 178
  mechanical, 28, 44, 133
  relational, 179
  statistical, 179

Extended phenotype, 167

Extracellular matrix (ECM), 253, 255, 256, 258, 269, 271, 276, 278

## F

Fibroblasts, 269, 278, 279

Fischer, K., 24

Fitness, 43, 135, 137, 138, 143, 149, 165, 166, 175, 199, 220, 234, 288, 298, 309

Function, 10, 15, 24, 26, 29, 31, 32, 36, 285-287, 289, 300, 318
  ecological, 285-322
  and functional domain, 203-205
  organizational, 291-310, 313-315, 317

## G

Galen, 24

Gauss, C.F., 35

Gemüt, 77

Genetic
  code, 138, 210
  determinism, 167
  program, 3, 264

Genocentrism, 1-4, 6, 225, 226, 228

Girtanner, C., 28

Glisson, F., 27
Goldstein, K., 44, 46, 48, 51, 52, 69

Gould, S.J., 128–130, 135, 223, 230

Grene, M., 44, 46, 48

Griesemer, J., 162

## H

Heredity, 165, 169, 172, 175, 188, 199, 202,
203, 212, 229, 259

principle of, 165, 169, 175

Hertwig, O., 29

Heterogeneity, 64, 71, 77, 206

Heuristic of collaboration, 223, 225, 231

Historicity, 15, 266, 268, 269, 280

Holism, 48, 50, 53

Hooke, R., 26

Hormones, 270, 278–280

Hypercycle, 170

## I

Individual, 15, 266

biological, 36, 120, 129, 141, 144, 175,
202, 208

Information, 264

Inheritance, 4, 128, 167, 168, 177, 191,
204, 219–236

extended, 4, 5, 219, 224–226, 230

Integration, 2, 3, 9–11, 30, 43, 52, 93, 95,
129, 146, 202–204, 224, 225,
234, 235, 253, 255, 258, 271,
280, 301, 302, 311

functional, 189, 296

Interdependence, 30, 31, 36, 189, 194, 226,
232, 233, 235, 265

In vitro, 188, 189, 271, 278–280

Items

abiotic, 314

of biodiversity, 314

## J

Judgement

broken, 63–65

determinative, 73, 81

reflective, 73, 81, 140

Jungius, J., 26

## K

Kant, I., 12, 16, 28–30, 32, 34, 37, 42–44, 47,
54, 60, 62, 69, 75, 77, 78, 80, 81,
129–141, 148, 149, 151, 192, 223, 226,
317, 318

Kauffman, S., 14, 36, 226, 250, 265

autocatalytic set, 170, 193, 235, 301, 311

edge of chaos, 170

work-constraint (W-C) cycle,
14, 36, 90, 164

## L

Lamarck, J.-B., 30

La Mettrie, J.O. de, 26

Laughlin, R.B., 117, 118

Leibniz, G.W., 27, 67

Levels, 1, 2, 14, 24, 30, 32, 33, 36, 37, 52,
128, 132, 137, 141–144, 146, 160,
173, 179–181, 190, 191, 193, 195,
197–199, 201–203, 205, 207,
209, 224, 248–251, 255, 258,
265, 268, 272

Lewontin, R.C., 42, 128–130, 135, 165, 166,
169, 170, 175, 223, 230

Life cycle, 139, 168–175, 180, 181, 233, 242,
244–249, 257, 259, 276

Lineages, 128, 166, 170, 174, 199, 201, 202,
206, 211, 212, 228, 233, 234, 242, 243,
251, 272

Linnaeus, C. von, 28

Living beings, 12, 24–32, 34–37,
42, 44, 50, 60, 68, 75, 98,
192, 210, 269, 318

Lloyd Morgan, C., 111, 113

Locke, J., 27

## M

Mammary glands, 265, 269, 278

Materiality of biological objects, 266

Mathematical modeling, 274, 275, 279, 280

Maturana, H., 90, 196, 250, 265

Mechanism, 12, 13, 24, 28, 29, 45, 47, 49, 53,
54, 130, 152, 160, 178, 191, 193,
205, 209, 210

Minimal metabolism, 204

Modularity analysis, 303–305, 307

Molecular reaction networks, 211

Moral

agents, 316, 317, 320, 321

patients, 316, 317, 321

value, 317

More, H., 26

Moreno, A., 35, 50, 53, 91, 250, 259, 291,
299, 315

Morphogenesis, 171, 265, 269–271,
273, 279

morphogenetic fields, 168, 179
Morphology, 35–37, 129, 244, 269, 270

Mossio, M., 36, 50, 91, 255, 291

Müller, J., 12

Multicellularity, 142, 143, 253, 260

N

Natural purposes, 29, 30, 43, 75, 129, 131–134, 140, 149, 150

Natural selection, 165–167, 288, 298, 307, 310

conditions for, 136, 165–167, 169, 170, 172, 175

units of, 121, 166, 168, 169, 171, 176, 181

Needful freedom, 164

Needham, J., 13, 28, 47, 48, 51

Neomechanism, 53

Newton, 10, 47

Newtonian paradigm, 48, 177, 178, 180, 181

Niche construction, 4, 5, 128, 148, 177, 221

Normativity, 15, 91, 98, 131, 179, 264, 266, 273, 285–287, 289, 293, 303, 307, 315, 316

O

Ontology, ontologization, 42, 43, 45, 47, 48, 50–52, 54, 86, 92, 133, 153, 161, 177, 265

Organic bodies, 24, 25, 30, 31

Organicism, 15, 41–54, 103–123, 264, 265

Organism, 2, 11, 24, 41, 127, 159, 189, 219, 242, 264

Organization, 1, 2, 29–32, 50, 77, 86–88, 105–107, 135, 162, 164, 165, 189, 194, 199, 201, 202, 206–210, 227, 243, 249–251, 253–255, 258, 266, 286, 289, 290, 315

levels of, 53, 87, 88, 161, 166, 167, 170, 173

narrow vs. wide, 87

vs. order, 24, 31, 33, 88, 99

principle of, 8, 11, 173, 174, 196, 227, 268, 275, 280, 308

3 degrees of explanatory centrality, 85

Organizational continuity, 162–165, 172–176, 181, 255

Origins of Life, 8, 191, 192, 195

“metabolism first” vs. “replication first” view, 4, 8, 189, 209

P

Pathology, disease, 267

Pattee, H.H., 35, 250

Perspectival realism, 161, 162, 166–169

Phenotypic plasticity, 4, 5, 128, 177

Physicalism, 112, 113, 115, 264

Piaget, J., 13, 31, 163, 250

Plessner, H., 32

Pluralism, 313, 320

Polanyi, M., 35

Porphyrios, 25

Prebiotic transitions, 212

Primordial biogenesis, 190, 191, 196, 197, 202, 205, 207, 210–212

Principle of biological inertia (default state), 268, 271, 275, 280

Process

ontology, 163, 168

structuralism, 168, 179

Protocell models, 8, 9, 189, 191–198

Proto-phylogeny, 201, 202

Q

Queller, D.C., 144, 145, 296

R

Rationality, 61, 68, 77, 79–81

Reciprocity, 29, 30, 61, 65–69, 71, 73, 77, 178

causal, 160, 171

Reductionism, 161, 264, 265

Reflexivity, 52, 65–69

Regulation, 33, 34, 90, 92–95, 98, 134, 190, 206–210, 212, 251, 252, 256, 258, 299

Replication, 167, 169–174, 209, 220, 221, 223–225, 227, 229–231, 234–236

Reproducer perspective, 169–173

Reproduction, 11, 26, 32–34, 36, 43, 89, 142–144, 165, 167, 171–177, 180, 191, 195, 197–202, 204, 205, 210, 211, 230, 234, 242–249, 251, 255, 257, 266

Reuleaux, F., 35

Rosen, R., 14

(M,R)-systems, 9, 43, 193

Roux, W., 37
S

Schiller, F., 25, 27, 31, 77

Sedimentation, 191, 199, 203, 204, 209

Self-organization, 11, 12, 24, 33, 43, 66, 75,
77, 133–135, 138–141, 149, 153, 164,
171, 172, 192–194, 206, 207, 226, 266

Self-referentiality, 30, 32, 36, 37

Sextus Empiricus, 25

Socioenvironmental crisis, 309

Soul, 24–28, 54

Stahl, G.E., 30

Stochastic corrector model, 170

Strassmann, J.E., 144, 145, 296

Stroma, 269–271, 279

Subject, 66, 82

Subjectivity, 47, 50, 51, 53, 54

Subject/predicate structure, 61, 68

Sustainability, 196, 212, 285, 287, 321

Systems biology, 6–8, 10, 43, 128, 189, 193,
195, 264

T

Teleology, finality, purposiveness, self-
directedness, 11, 15, 30, 32, 93,
285–287, 289, 293, 295, 303, 309,
315, 319

and anticipative learning, 88, 93, 94, 96, 98

and explanation, 46, 179, 181, 266,
286, 287

intrinsic, 227, 251, 266, 286, 287, 289,
295, 307, 316, 318–321

and self-determination, 11, 164, 165,
286, 319

Teleonomy, 176, 279

Theoretical Biology Club, 13

Theoretical principles, 9, 15, 227, 265, 268,
274, 275

Thermodynamic
openness, 11, 13, 14
predicament, 164

Thompson, E., 47, 50, 53, 69

3D culture, 271, 273, 275

V

Valéry, P., 34

Value
instrumental, 295, 317
intrinsic, 318, 321
moral, 316, 318

Varela, F.J., 15, 33, 47, 50, 90, 196, 250, 265

Variation, 11
heritable, 128, 165, 220, 225, 228, 230,
231, 236
principle of, 165, 169, 171, 268, 274,
280, 308

Vitalism, 13
attitudinal, 62, 67, 69, 78–81
heuristic, 69
substantival, 69

Vleeschauwer, H. de, 65

Voltaire, 28

von Neumann, J., 32

W

Waddington, C., 13

Weiss, P., 13

Whole, wholes, 46, 141

Woodger, J.H., 13, 50, 60, 66, 68,
81, 106, 120