8. Which Phenomena Are Function-Based Living Individuals?
The functional-network-based definition of a living individual encompasses the cybernetic definition proposed previously (Korzeniewski 2001, 2005, 2023). Therefore, phenomena that were counted as cybernetic individuals also belong to function-based individuals. However, the justification of their classification is frequently significantly different and the functional definition delivers significantly stronger conceptual tools to deal with this problem than the cybernetic definition.
First, all processes traditionally regarded as inanimate phenomena, such as stones falling in the gravitational field, crystals growing in a saturated solution of salt or “flowers” “painted” by frost on window glasses certainly do not constitute (elements of) biological systems in general, and function-based individuals in particular. They are not purposeful, shaped by biological evolution, directed on the realization of the superior goal of self-sustaining and proliferation. Some processes, like convective currents belonging to dissipative structures, are dynamic processes organized in space and time, sustained by a ceaseless circular flow of matter, dissipation of energy and production of entropy. However, they lack any operational intentionality as they are not directed on any ultimate task. They are not (composed of) Ps, Ss, Fs, R(s) and D(s).
Some devices produced artificially by humans, for example robots equipped with computers, exhibit a set of functions and realize concrete tasks. Several functions occur within and are carried out by them. They comprise passive purposeful processes (e.g., movements), signal transduction (e.g., electrical), negative feedbacks (e.g., responsible for precise control of operations or navigation in the surroundings), re-writing processes (e.g., mechanistic information copying by computers) and decoding (arbitrary transformation of information from one form into another using available hardware and software by computers, for instance during constructing of complex devices on the basis of sets of instructions). However, these functions and tasks of human-produced devices are not “their own”. On the contrary, they are imposed to them by their human creators to whom these devices serve. They do not reproduce by themselves (copy themselves), possess any mechanisms directed on this purpose, and can be only produced in human factories in a certain, frequently large number of copies. Certainly, they are not self-dependent, self-sufficient in and self-directed on the production of themselves. Therefore, they are by no means artificial function-based living individuals.
However, there exist numerous biological phenomena that are “alive” in this sense that they are complex sub-systems of living systems or are composed of living individuals (of the same or different species), but they are not function-based living individuals themselves. There also exist systems that have only a limited autonomy as individuals and therefore are only individuals to a certain degree and not fully-fledged function-based living individuals (their IA < 1). Let us consider some doubtful cases in order.
Viruses are frequently regarded as situated on the border of life and non-life. Their infectious particles outside the host cell, virions, have the form of a nucleic acid molecule (one- or two-stranded, linear or circular RNA or DNA) encapsulated in a protein capsid composed of single protein molecules: capsomers (Harris and Hill 2021). They exhibit no “vital functions”: metabolism, movement, growth or replication. They execute no Ps, Ss, Fs, R(s) and D(s). Are they therefore “dead”? Not at all. They resemble e.g., bacteria spores produced in unfavorable (hostile) conditions for the purpose of survival that are completely dehydrated (devoid of water) and carry out no metabolism and other functions. However, as discussed above, it is not just one stage, but the whole life cycle of a given organism that constitutes the function-based living individual. When a virion enters the host cell, the situation changes dramatically. It takes over the control of the cell functioning, re-directs its metabolism and genetic apparatus (all its Ps, Ss, Fs, R(s) and D(s)) and forces it, instead to perform normal functions and maintain homeostasis, to produce maximal possible amount of copies of the virus particles (virons), which dramatically disturbs the cell functioning by depriving it of building substances and energy, leading eventually to its death (not all viruses are lethal for the infected cell). The virus is very efficient in the realization of this task. These functions of the host cell captured by the virus and imposed by itself involve p-processes (e.g., synthesis of organic compounds being substrates for new virions production), signals and feedbacks (taking over and controlling the cell functioning, but also many of the existing regulatory mechanism of the cell metabolism), re-writing processes (i.e. copying the virus nucleic acid) and decoding processes (e.g., synthesis of proteins encoded in this nucleic acid such as capsomers or reverse transcriptase in retroviruses) (anyway, viruses also use numerous proteins already synthesized by the host cell). Of course, all these functions are executed by the host cell components (especially enzymes) or by a few proteins synthesized by these enzymes on the matrix of the virus nucleic acid. However, the essence of being a parasite is to get rid of as many vital functions of its own as possible and to “cede” these functions to the host organism. Viruses realize this strategy to an extreme extent. While isolate virions outside host cells do not seem “alive”, viruses “dipped” in the functional network of the host cell switched by the virus to its own proliferation certainly behave as fully-fledged living individuals. Additionally, as the virus RNA or DNA undergoes mutations, viruses are subject to normal natural selection and Darwinian evolution. Therefore, viruses are certainly function-based living individuals, although with a very low value of IA, much more close to 0 than 1.
Viroids are even simpler than viruses, as their naked circular single (but containing many intra-molecular nitrogenous base-pairing regions) RNA strand contains only a few hundreds of nucleotides and they are devoid of the protein capsid (Navarro et al. 2021). However, essentially all the reasoning concerning the status of viruses as function-based living individuals refers also to viroids. When they enter into the host angiosperm plant cell, its functioning is re-directed and it executes all the functions necessary for the viroid RNA to replicate: Ps, Ss, Fs, R(s) and D(s) (of course, most of them are functions that normally occur in non-infected cells). To be sure, viroid RNA does not code for any protein. However, this does not mean that decoding is not involved in the viroid proliferation. All the necessary proteins are, before infection or under the viroid “command”, synthesized by the cell’s genetic apparatus. For example, the viroid RNA is replicated by enzyme RNA polymerase II, which in a “normal” (not infected) cell catalyzes transcription of DNA to mRNA. Therefore, viroids are also, like viruses, function-based living individuals, although characterized by an extremely low, even lower than in viruses, value of IA.
Depending on the particular case and conditions, plasmids (Kado 1998; Brown 2012) can be regarded as either mutualistic symbionts or parasites of bacteria and some simple eukaryotes. They are small extrachromosomal circular DNA molecules present in one or several copies in the host cell. The proliferate independently (they are replicons) and code for they own system of replication. They also code for proteins controlling their separation to descendent cells during the host cell division. They can contain the addiction system that kills the host cells that have not inherited a copy of the plasmid during cell division. They can transfer themselves between host cells during the process of conjunction. They are not essential for the host functioning and survival under normal conditions. However, they can be very helpful in special circumstances, as they can condition the resistance to antibiotics, ability to decompose poisons, induce virulence or code for a system being a substitute of sexual reproduction: when transferring themselves (their copies) from a cell to a cell, they can carry with them fragments of the host genomes and thus participate in the horizontal gene transfer. Generally, they are equipped with a rich network of functions, including Ps, Ss, Fs, R(s) and D(s), both their own and that of the host cell. Therefore, they certainly should be counted among function-based living individuals with a relatively high IA value, undoubtedly higher than in viruses and viroids.
Transposons (transposable elements (TE), jumping genes) (Pray 2008; Brown 2012; Futuyma 2013), counted among parasitic DNA (Doolittle and Sapienza 1980; Orgel and Crick 1980), are mobile genetic elements – segments of DNA of eukaryotic chromosomes that can change their position within the genome. There are distinguished two apparently quite similar classes of transposons. Class I transposons (retrotransposons) pass through the RNA stage during their “life cycle”. They encode a few proteins (D), including reverse transcriptase, which catalyzes synthesis of a DNA strand on the matrix of an RNA strand (R). The regular cell DNA-dependent RNA polymerase carries out the transcription of the retrotransposon DNA to RNA (R). Next reverse transcriptase rewrites this RNA to a single DNA strand (R), that is next doubled by DNA-dependent DNA polymerase (R). The newly-synthesized double DNA segment of the retrotransposon is then inbuilt in a new place in the “host” genome. In this way, the number of the transposon copies can be substantially increased. Class II transposons (DNA transposons) do not pass through the RNA stage. They code for enzyme transposase (D)responsible for insertion and excision of DNA transposon to/from the “host” DNA. The whole process of transposition consists in just relocation of the transposon from one place to another. The number of transposons does not increase – they do not proliferate, but just relocate themselves. Both classes of transposons deprive the host cell of building substances and energy and unnecessarily (to our knowledge) occupy space in chromosomal DNA. Of course the whole process of transposition of Class I and II transposons is involved the functional network (Ps, Ss, Fs, R(s) and D(s)) of the “host” cell with a minor contribution of the transposons own mechanisms. However, retrotransposons do proliferate and they are purposefully directed on this task, albeit only within the genome of the “host”, while DNA transposons just travel from one place in the genome to another, although they are evidently purposefully directed on doing so. Therefore, while retrotransposons are function-based living individuals, albeit with a very low IA, lower even than in viruses, then DNA transposons are not. Therefore, the purposeful transposition of the latter remains a puzzle unless it is somehow supported by the “host”. Anyway, the “environment” of the former in which they can spread remains very limited: it is just the genome of their “host” (or “carrier”), and they can “infect” only the haploid genome coming from the other parent (its gamete) in the fertilized egg cell (zygote). The genetic record of retrotransposons can mutate and evolve, as various transposons can compete for the limited “environment capacity” in the genome, which is a hallmark of a living individual.
Supernumerary chromosomes B (Jones 1975; Bougourd and Jones 1997), another type of parasitic DNA, are unfunctional degenerated chromosomes that nevertheless are present in genomes of many eukaryotic species. The individuals devoid of chromosomes B do equally well, if not better, than their “carriers”. It seems that organisms try to get rid of chromosomes B and that the latter have purposeful mechanisms counter-acting this (Nur et al. 1988). They most probably evolved from “normal” functional chromosomes. Of course, chromosomes B are sustained and replicated with the aid of the whole metabolism and genetic apparatus (all Ps, Ss, Fs, R(s) and D(s)) of the cell containing them. However, they cannot proliferate within the cell. In this aspect they resemble Class II (DNA) transposons. In such a case, are they not living individuals at all? The situation is not so simple, as they are replicated by the cell genetic apparatus during cell division, including meiosis. If they are able to get preferably to gametes, especially the egg cell, they can “infect” the fertilized egg cell containing half of the genome coming from another parent that might not to contain B chromosomes of a given type. In such a way, chromosomes B could propagate and thus deserve classifying as function-based living individuals, although only to a very rudimentary extent with a very low IA. Furthermore, if a given type of DNA transposon (Class II) can also be preferably transferred to gametes, it should regain the status of a function-based (barely) living individual. As it seems, the fact of being or not a function-based individual is not such a clear-cut distinction as could be expected.
Prions (Prusiner 1998) are pathogenic misfolded protein molecules that cause neurodegenerative diseases of the brain in mammals (including humans). The normal (correctly folded) molecules of the prion protein (PrP), possessing α-helix structure, are encoded in the animal genome, synthesized in the brain and other organs and fulfil useful, still not fully understood, function in the organism. The misfolding or transition to the pathological form of a PrP molecule, resulting in the β-sheet structure, can be spontaneous (rarely) or induced by an already misfolded PrP molecule. Such molecules can come from consumed brain tissue of another animal (or human in the case of New Guinea cannibals). Each pathogenic molecule can induce misfolding of several normal molecules, which can in turn induce misfolding of subsequent molecules. In the result of this, the number of misfolded molecules increases exponentially and a positive feedback appears, analogous e.g. to a stony avalanche or bacteria proliferation on laboratory medium. Like viroids, prions are single molecules of an organic linear hetero-polymer, this time protein. Like all proteins synthesized by a living cell, its production is involved in a rich network of functions (Ps, Ss, Fs, R(s) and D(s)). They even exhibit a rudimentary form of replication: pathological molecules proliferate at the cost of normal molecules. Therefore, are they function-based living individuals? Of course not. They constitute a case of a spontaneous living system misfunctioning, something resembling to some extent auto-allergy. They are equipped with no functions/mechanisms purposefully supporting (directed on) their reproduction (IA = 0). They cannot evolve in order to increase their “virulence”, as their identity is encoded in the cell DNA. They just constitute a kind on an “evolutionary trap”.
Cancer cells (Papaccio et al. 2017) originate in the result of tumorous transformation caused by somatic mutations in normal cells. They proliferate in the place where the transformation took place and possibly migrate to other organs and there also proliferate forming metastases. Eventually, they die together with the death of the host organism, frequently caused by them. The divisions of normal cells are strictly controlled by the organism. Mutations cause cancer cells to escape this control and increase their efficiency in proliferation and migration. Traditionally, cancer cells are not regarded as living individuals, perhaps partly because they are still almost identical genetically with their “carrier”. However, the tumorous transformation of a normal cell constitutes breaking out from the superior purpose of the whole organism and establishing its own superior purpose, that is its own survival, reproduction and spreading in the “environment” of the host organism. Cancer cells are subject to normal Darwinian selection and evolution. Only those cells survive and proliferate that develop (due to mutations) numerous adaptations involving purposeful functions of their own (comprising Ps, Ss, Fs, R(s) and D(s)) that support and are directed on their survival, replication and spreading within the host organism. These functions include:
insensitivity to the organism’s cell proliferation control;
resistance to the immunological system and other defensive mechanisms of the host organism;
adaptation of metabolism to lowered oxygen levels, for instance anaerobic (glycolytic) ATP production; the neoplastic tumor has limited vascularization and thus blood flow;
stimulation of angiogenesis (formation of blood vessels) in the neoplastic tumor;
resistance to the induction of apoptosis (programed cell death);
maximization of the rate of cancer cells divisions;
maximization of the rate of cancer cells migration and metastases formation;
competition with other cancer cells, with somewhat different genetic identity, for the limited capacity of the “living space” within the host organism.
Therefore, in the “environment” of the host organism cancer cells behave as regular fully-fledged living individuals. From normal parasites, they differ mainly by the genetic near-identity with the host and impossibility to infect other hosts. However, there are known examples of cancers that can be transferred from one host individual to another, such as dog venereal sarcoma that is transferred through sexual contact in dogs (Barski and Corneferd-Jensen 1966) or facial tumor in Tasmanian devils (Woods et al. 2015), where mutated tetraploid lymphocytes spread to other devils by bites during fights for sexual mate or territory. Generally, cancer cells decidedly constitute function-based living individuals with quite a high value of IA.
In some cases two initially completely independent function-based individuals (IA = 1) start to cooperate and support each other through offering to the partner some functions the partner does not possess and using some functions of the partner that the individual in question is not equipped with. Both partners benefit from such cooperation provided that the advantages prevail over the costs. Mutualistic symbionts are individuals of (very) different species that entered into such a relation called mutualism. It can be facultative, when the partners can, but do not have to cooperate (they can survive and reproduce, although less successfully, in separation), or obligatory, when the partners are absolutely necessary for each other in order to survive and/or reproduce. Examples of facultative symbiosis are cleaner fish and their “clients” – bigger fish (e.g., sharks), whose teeth and skin they clean. The cleaners get food and the cleaned – more healthy teeth and skin, but they can manage alone. In this case, IA of both cleaner fish and their clients is decidedly close to 1. Ants of many species protect and in fact “culture” aphids that in return feed them secreting honeydew. However, both ants and aphids can survive without their partners (ants would have to look for another source of food and aphids would be less protected). Therefore, for both of them 0 < IA < 1. In most cases of lichens, conglomerates of the alga and fungus components (fungi supply water and mineral substances from the substrate, while algae – organic products of photosynthesis), the partners cannot live alone under natural conditions. However, in some cases they can, and they can be cultivated separately in laboratory conditions. Therefore, IA of algae and fungi is closer to 0 than in the previous case, or simply reaches 0. There are many examples of obligatory mutualism, partnership “for life and death”. They include mycorrhiza, that is cooperation between a fungus that fulfill the role of root hair and takes up water and minerals from the soil, and a vascular plant that provides the fungus with organic products of photosynthesis. Flowering plants are fertilized by various insects, birds or bats (fulfilling the role of pollinators) and in return offer sweet nectar and pollen as food, using their colorful flowers as a lure. Termites cannot digest by themselves the dead wood they feed on and they culture for this purpose protozoans or bacteria in their guts: termites get easily assimilable organic compounds, while protozoans/bacteria – raw food and place to live. As the close relationship between obligatory mutualistic symbionts tightens, their value of IA approaches 0. Although the partners in symbiosis retain much of its autonomy, they cannot survive and reproduce independently. If there are still some genetic/evolutionary conflicts between the partners, as they still have their own reproductive systems, IA is slightly greater than zero. If not, it reaches zero. The degree of individuality is not the same what autonomy. For example, the living cell or liver in a complex animal organism possess quite a great degree of autonomy, but they have no individuality (see below).
Mitochondria, called the power plants of eukaryotic cells that deliver most of energy in the form of ATP through oxidizing organic compounds (respiratory substrates) using oxygen, are descendants of once free-living α-proteobacteria (IA = 1) (Reece et al. 2011). They entered into endo-symbiosis (living within) with ancestors of eukaryotic cells: Asgard archaea. Apart from other functions, mitochondria deliver ATP to the cytosol of the eukaryotic cell, receiving from it respiratory substrates as well as most of proteins, carbohydrates and lipids they are built of. Most of their genes have been transferred to the nucleus or lost and presently they are significantly simplified in relation to their ancestors. Nevertheless, they have still preserved some of their reproductive autonomy, although their number is strictly controlled by the cell. On the other hand, mitochondria never originate de novo but only through the division of maternal mitochondria. They contain several copies of their own circular DNA, code for some proteins, for example some subunits of the oxidative phosphorylation complexes, for their own tRNA and rRNA, possess small ribosomes much more resembling bacterial ribosomes than that situated just outside them in the cytosol, are encompassed by two lipid-protein membranes. The mitochondrial DNA still can mutate, and therefore particular mitochondria can possess their own identity and potentially care about their own (evolutionary) interests. For example, in plants, there can occur a genetic conflict between mitochondria and the “cytosolic rest” of the plant. Mitochondria are transferred only through female gametes (egg cells) and therefore a mitochondrial gene cms prevents production of male sex organs in the flower: the stamens. However, the plant activates its own nuclear gene r that suppresses the action of gene cms (Futuyma 2005). On the other hand, the mitochondrial DNA in animals, especially mammals, is much smaller than in plants and contains no such “conflicting” genes. It is not clear whether mutated and thus difunctional mitochondrial can be somehow eliminated by host cells – certainly such mutations cause mitochondrial myopathies (MMs). Taking into account all these facts, mitochondria can be treated either as rudimentary individuals (IA very close to 0), descendants of fully-fledged free-living individuals, or as objects that have already completely got rid of their individuality (IA = 0).
Hydrogenosomes (Yarlett and Hackstein 2005) evolved from mitochondria and are organelles that produce ATP in the process of fermentation in certain primitive protists living in strictly anaerobic (devoid of oxygen) conditions. They are encompassed by a double protein-lipid membrane, but devoid of their own DNA and genetic apparatus. Therefore, in this case the fusion of initially two separate individuals into one has ultimately proceeded to the very end. Hydrogenosomal IA = 0.
Chloroplasts (Reece et al. 2011) are to some extent analogous to mitochondria. They are organelles carrying out photosynthesis in plants and are (descendants of) bacterial endosymbionts, this time cyanobacteria that settled in the ancestors of plants. They have a significantly greater DNA encoding more proteins and more complex structure than mitochondria, their own genetic apparatus and lipid-protein membranes. Therefore, they are rudimentary function-based individuals with IA close to 0, but probably slightly greater, than in mitochondria. However, it cannot be excluded that their IA = 0.
Colonial coelenterates, bryozoans and tunicates (Reece 2011) are composed of “quasi-individuals” or polyps/zooids of the same or different types (e.g., feeding, defensive, reproductive, locomotory). Genetically, they are colonies of clones budding from one larva or ancestral zooid developed from the zygote. The polyps/zooids have connected digestive systems – in fact the share one common gastrovascular cavity/gut – and they exchange food. They are to some extent physiologically (functionally) integrated and exhibit cooperation. The whole colony controls the budding and growth of particular (types of) polyps/zooids, depending on conditions. Therefore, in the case of functional (and, of course, structural) differentiation of polyps/zooids (into sexual and fulfilling other functions), the whole colony constitutes a function-based living individual, as only the whole colony can survive and reproduce. However, even in the case of one type of polyps/zooids that can reproduce, they are all genetically identical and can be regarded as a single function-based individual. On the other, if some mutations occur during the colony growth by budding, different zooids start to differ genetically and therefore to possess different identities they tend to proliferate. Therefore, in this case such mutated zooids become to a tiny extent separate function-based individuals. Anyway, this concerns in fact all organisms that do not possess separated generative (germ) cell lines and thus are in fact well-integrated colonies of somatic groups of cells, for instance trees, where “mutated” branches (somatic groups of cells) with “mutated” flowers produce gametes with a modified identity. Summing up, this is the whole colony of coelenterates or bryozoans that constitutes the function-based individual (IA close to 1), while individual zooids have zero or very low level of individuality (IA equal or very close to 0).
Another interesting case is slime molds (mycetozoans) (Maynard Smith and Szathmary 1995), distant relatives of fungi. During most of their life cycle they have the form of unicellular amoeboids (amoeba-like cells) spread over some area and feeding independently. However, in the case of nutrients deficit, the amoeboids approach each other attracted by chemical signals (chemotaxis) and congregate into multi-cellular plasmodium (slug). After some time the plasmodium forms a sporangium (fruiting body). The chamber of the sporangium is situated at the top of a stalk and contains spores. When the wall of the sporangium breaks, the spores are disseminated. In hospitable conditions (plentiful food) they develop into amoeboids that feed and proliferate and the cycle closes. It is uncertain whether particular amoeboids or the whole plasmodium constitute the function-based living individual. This depends on the genetic identicalness of amoeboids in the plasmodium. If the whole local population of amoeboids comes from spores of one plasmodium, the whole population of cloned amoeboids simply constitutes an “individual in dispersion”, which undergoes consolidation into plasmodium to produce and disseminate spores. If, however, a few genetically different amoeboid groups enter into the plasmodium composition, either because they come from spores produced by different parental plasmodia or they originated in the result of sexual reproduction that facultatively occurs in slime molds, an interesting case of an intra-species reproductive symbiosis (mutualism) takes place. However, this automatically creates a possibility of “reproductive parasitism”, where one of the genetically different groups of amoeboids possesses a mutation that would cause all amoeboids from this group to transform themselves selfishly into spores instead of honestly co-creating the stalk and sporangium walls with other groups of amoeboids. Of course, the mutated amoeboids will transfer more efficiently their identity (genes) to the progeny. Therefore, perhaps appropriate mechanisms preventing such a situation and suppressing the selfish strategy develop. Generally, it is likely that the plasmodium constitutes a kind if intra-species mutualistic symbiont composed of partial individuals (genetically identical groups of amoeboids) cooperating for the common good, with IA between 0 and 1, varying significantly depending on the stage of their life cycle, as a given amoeboid can potentially enter into co-operation with different other amoeboids.
Eusocial insects (wasps, bees, ants and termites) (Wilson 1971) and one rodent species (the naked mole) form colonies containing sexual forms and sterile (unable to reproduce or with suppressed reproduction) forms. The former can be one or more queens (fertile females) in hymenopterans (the males die after copulation) or a queen and a king (a fertile male) in termites and hairless moles. The latter may be workers (females in hymenopterans, females and/or males in termites and naked moles), frequently of a few kinds of different sizes and morphology (and, of course, fulfilling different functions) or soldiers (females in ants, females and/or males in termites). Insects of one type form casts. The role of queens and males is in principle limited to reproduction. The sterile casts realize all other tasks: taking care of queens (and kings), eggs and larvae, feeding them, acquiring food, building the nest, controlling its internal conditions, cultivating “fungi gardens”, defending the colony and so on.
Individual insects, both reproducing and sterile, although each equipped with a reach network of functions (almost all Ps, Ss, Fs, R(s) and D(s) present in solitary insects constituting living individuals), are not self-sufficient in the realization of the superior goal of the function-based individual: self-copying (propagation of own identity), requiring self-dependent sustaining of own existence (at least to the reproduction period) and reproduction. Obviously, sterile insects cannot reproduce and sexual insects (and their progeny) cannot sustain alone their functioning and survive without the help of sterile insects. Sexual and sterile forms must cooperate in order to realize the superior goal of a certain greater whole – the entire colony, its survival and reproduction (propagation of its identity). Particular insects in a colony are interconnected by a reach network of functions involving coordinated cooperation, communication and division of labor. These functions include:
► exchange of food between fed workers and hungry sexual forms, larvae, workers and soldiers (Ps, Fs);
► cooperation in building and repair of the nest involving communication and coordinated actions (Ps, Ss, Fs);
► maintaining homeostasis, involving regulation of the temperature and humidity in the nest, removing dead insects, scraps and debris (mostly Fs);
► cultivation of fungi gardens (Ps, Ss, Fs);
► marking odor traces or carrying out waggle dances indicating newly discovered sources of food (Ss);
► releasing chemical alarm substances in situations of danger (Ss);
► coordinated defense of the nest or column of insects outside the nest (Ps, Ss, Fs);
► common expeditions for food, forming floating islands or hanging bridges from coupled bodies of ants (Ps, Ss, Fs);
► regulation of the numerical strength of particular casts by pheromones or appropriate feeding of larvae (Fs).
Therefore, of course this is the whole colony and not particular sexual or sterile insects that constitute the function-based living individual. In most cases (apart from e.g., primitive colonial wasps) the value of IA for both reproductive and sterile insects is zero. It is frequently argued that sterile workers sacrifice themselves for the queen in the frame of the so called “kin altruism”, as they are very closely related to her and between them. However, in most cases this reasoning is wrong. Leaving apart the cases where a queen is fertilized by many males (which dramatically lowers the statistical kinship of particular workers) or many not-related queens co-exist within a single nest/colony and are supported by the same workers, the sterile insects do not choose their fate, but it is determined genetically, by larvae feeding or pheromones – they “have no choice”. While it is likely that the kin altruism played an important role in eusocial insect evolution, in developed insect societies it can be neglected (see Korzeniewski 2023 for discussion). From the formal point of view, eusocial insect colonies resemble multicellular organisms. Both systems are composed of sexual/reproductive and sterile/somatic elements: insects/cells and casts/organs, interconnected by a rich causally-closed network of functions directed on the proliferation of the identity of the whole system. Multicellular organisms evolved from unicellular ones, while eusocial insect colonies from solitary insects. In both cases the elements ceded their independence and autonomy to the entire system. In both cases, the “selfish” proliferation of the non-reproducing component elements of the whole: sterile insects/somatic cells is strongly suppressed by the whole (cancer cells or functional workers trying to lay eggs are rare exceptions from this rule). The main difference between them is that cells in multicellular organisms are integrated both structurally and functionally, while insects in insect colonies – mostly functionally (Korzeniewski 2023). Therefore, eusocial insect colonies even more deserve the name “functional individuals”.