Spatial Organization
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Spatial organization can be observed when components of an
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them und ...
or biological group are arranged non-randomly in space. Abiotic patterns, such as the ripple formations in
sand dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
or the oscillating wave patterns of the
Belousov–Zhabotinsky reaction A Belousov–Zhabotinsky reaction, or BZ reaction, is one of a class of reactions that serve as a classical example of non-equilibrium thermodynamics, resulting in the establishment of a nonlinear chemical oscillator. The only common element in ...
emerge after thousands of particles interact millions of times. On the other hand, individuals in biological groups may be arranged non-randomly due to selfish behavior, dominance interactions, or cooperative behavior.
W. D. Hamilton William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century. Hamilton became known for his theoretical work expounding a ...
(1971) proposed that in a non-related "
herd A herd is a social group of certain animals of the same species, either wild or domestic. The form of collective animal behavior associated with this is called '' herding''. These animals are known as gregarious animals. The term ''herd'' ...
" of animals, the spatial organization is likely a result of the selfish interests of individuals trying to acquire food or avoid
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
. On the other hand, spatial arrangements have also been observed among highly related members of
eusocial Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
groups, suggesting that the arrangement of individuals may provide advantages for the
group A group is a number of persons or things that are located, gathered, or classed together. Groups of people * Cultural group, a group whose members share the same cultural identity * Ethnic group, a group whose members share the same ethnic iden ...
. Spatial organization can be defined as how something is organized internally as well as externally concerning its surroundings. It can be viewed both on a large or small scale and may also include in its discussion, the factors and conditions that lead to a specific organization pattern. In the frame of biology this may include the abiotic and biotic factors that a population is immersed in. In the frame of civilization, spatial organization may include if an area is rural, suburban, or urban. Spatial organization can even include how one manages and organizes their lawn to meet social norms.


In eusocial insects

Individuals in a
social insect Eusociality (Ancient Greek, Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative Offspring, brood care (including care of offspring from other individuals), ove ...
colony can be spatially organized, or arranged non-randomly inside the nest. These miniature territories, or spatial fidelity zones have been described in
honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the ...
s (''Apis mellifera''),
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s (''Odontomachus brunneus''; ''
Temnothorax albipennis ''Temnothorax albipennis'', the rock ant is a species of small ant in the subfamily Myrmicinae. It occurs in Europe and builds simple nests in rock crevices. Description This species has the typical ant body pattern of head, mesosoma and metas ...
''; ''Pheidole dentata''), and
paper wasp Paper wasps are a type of Eusociality, social vespid wasps. The term is typically used to refer to members of the Vespidae, vespid subfamily Polistinae, though it often colloquially includes members of the subfamilies Vespinae (hornets and yel ...
s (''Polistes dominulus'', ''Ropalidia revolutionalis''). While residing in these zones, workers perform the task appropriate to the area they reside. For example, individuals that remain in the center of an ant nest are more likely to feed larvae, whereas individuals found at the periphery of the nest are more likely to forage.
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, ecologist, and entomologist known for developing the field of sociobiology. Born in Alabama, Wilson found an early interest in nature and frequ ...
proposed that by remaining in small, non-random areas inside the nest, the distance an individual moves between tasks may be minimized, and overall colony efficiency would increase.


In the nest


Foraging-for-work

There are a variety of ways in which individuals can divide space inside a nest. According to the "foraging-for-work" hypothesis, adult workers begin performing tasks in the area of the nest where they emerged, and gradually move towards the periphery of the nest as demands to perform particular tasks change. This hypothesis is based on two observations: "(1) that there is spatial structure in the layout of tasks in social insect colonies and (2) that workers first become adults in or around the center of the nest". Individuals can remain in an area for an extended period, as long as tasks need to be performed there. Over time, an individual's zone may shift as tasks are accomplished and workers search for other areas where tasks need to be performed.
Honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the ...
s, for example, begin their adult life caring for brood located in the area near where they emerged (i.e. nurse bees). Eventually, workers move away from the brood rearing area and begin to perform other tasks, such as food storage, guarding, or foraging.


Dominance hierarchy

Space inside the nest may also be divided as a result of dominance interactions. For example, in
paper wasp Paper wasps are a type of Eusociality, social vespid wasps. The term is typically used to refer to members of the Vespidae, vespid subfamily Polistinae, though it often colloquially includes members of the subfamilies Vespinae (hornets and yel ...
colonies, a single inseminated queen may find (initiate) a colony after waking up from hibernation (overwintering). However, it is common in many species that multiple inseminated females join these foundresses instead of founding their own nest. When multiple inseminated females found a colony together, the colony grows quickly, yet only one individual will become the primary egg-layer. Through a series of dominance interactions, the most aggressive wasp will emerge as the dominant individual and will become the primary egg-layer for the group (the prime role for ensuring your genes are passed on to subsequent generations), whereas the remaining subordinate wasps will perform other tasks, such as nest construction or foraging. There is evidence that these dominance interactions affect the spatial zones individuals occupy as well. In paper wasps (''Ropalidia revolutionalis''), as well as in the ant species ''Odontomachus brunneus'', dominant individuals are more likely to reside in the central areas of the nest, where they take care of the brood, while the subordinate individuals are pushed towards the edge, where they are more likely to forage. It is unknown whether division of space or establishment of dominance occurs first and if the other is a result of it.


Outside the nest

There is also evidence that foragers, which are the insects that leave the nest to collect the valuable resources for the developing colony, can divide space outside the nest. Makino & Sakai showed that
bumble bee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related genera ...
foragers maintain foraging zones in flower patches, which means that bees consistently return to the same areas within a patch and there is little overlap between individuals. These zones can expand and contract when neighboring foragers are removed or introduced, respectively. By dividing foraging patches into miniature ‘foraging territories’, individuals can maximize the number of flowers visited with minimal interruptions or competition between foragers. These ‘foraging territories’ divided among individuals from the same colony are the result of
self-organization Self-organization, also called spontaneous order in the social sciences, is a process where some form of overall order and disorder, order arises from local interactions between parts of an initially disordered system. The process can be spont ...
among the foragers; that is, no lead forager is dictating where the bees will forage. Instead, the maintenance of these foraging zones is due to simple rules followed by each individual forager. Studies to determine these “rules” are an important area of research in
computer science Computer science is the study of computation, information, and automation. Computer science spans Theoretical computer science, theoretical disciplines (such as algorithms, theory of computation, and information theory) to Applied science, ...
, basic
biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
,
behavioral ecology Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for ethology, animal behavior due to ecology, ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined Tinbergen's f ...
, and mathematic modeling.


As an emergent property of a self-organized system

The
self-organization Self-organization, also called spontaneous order in the social sciences, is a process where some form of overall order and disorder, order arises from local interactions between parts of an initially disordered system. The process can be spont ...
observed in foraging territories is a microcosm for the self-organization seen in the entire
colony A colony is a territory subject to a form of foreign rule, which rules the territory and its indigenous peoples separated from the foreign rulers, the colonizer, and their ''metropole'' (or "mother country"). This separated rule was often orga ...
. Spatial organization observed across social insect colonies can be considered an emergent property of a self-organized
complex system A complex system is a system composed of many components that may interact with one another. Examples of complex systems are Earth's global climate, organisms, the human brain, infrastructure such as power grid, transportation or communication sy ...
. It is self-organized because no leader is dictating where each individual will reside, nor which task an individual will perform once they get there. Instead, zones may be a by-product of division of labor, whereby individuals end up in a particular location for a period based on the task they perform, or dominance interactions, whereby dominant individuals are granted access to the most desirable places inside the nest. Spatial patterns exhibited by individuals of social insect colonies are not obvious, because it is difficult to observe and differentiate among individuals inside a nest cavity or flying across a foraging patch. However, when careful attention is given to the individual worker, the spatial organization of workers in the nest becomes apparent.


See also

*
Cellular automaton A cellular automaton (pl. cellular automata, abbrev. CA) is a discrete model of computation studied in automata theory. Cellular automata are also called cellular spaces, tessellation automata, homogeneous structures, cellular structures, tessel ...


References

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