CHAPTER 1: HOW

Holistic approach and Systems’ thinking view of organisms and evolutionary processes

The methodology of Systems’ thinking is of understanding of a system by examining the linkages and interactions between the components that comprise the entirety of that defined system. (Summarized from wiki) A system may be defined in general as a set of interrelated or interacting elements. This definition accommodates both passive structures and active structures. In biology, a living organism is seen as a set of organs, muscles etc. that interact in processes to sustain the organism. Each cell is seen as a collection of organelles that interact in processes to sustain both the cell and the wider organism.

The concept of a system:
  • A system is composed of parts
  • A system is other than the sum of its parts
  • All the parts of a system must be related (directly or indirectly), else there are really two or more distinct systems
  • A system is encapsulated (has a boundary)
  • A system can be nested inside another system
  • A system can overlap with another system
  • A system is bounded in time, but may be intermittently operational
  • A system is bounded in space, though the parts are not necessarily co-located
  • A system receives input from, and sends output into, the wider environment
  • A system consists of processes that transform inputs into outputs
  • A system is autonomous in fulfilling its purpose
  • A system is a dynamic and complex whole, interacting as a structured functional unit circuit.
  • Energy, material and information flow among the different elements that compose a system.
  • A system is a community within an environment.
  • Energy, material, and information flow from and to the surrounding environment via semi-permeable membranes or boundaries that may include negotiable limits.
  • Systems are often composed of entities that seek equilibrium but can exhibit patterns, cycling, oscillation, randomness, or chaos, or exponential behavior.
  • A holistic system is any set (group) of interdependent or temporally interacting parts. Parts are generally systems themselves and composed of other parts, just as systems are generally parts (or “holons”) of other systems.
The systems thinking approach incorporates several tenets:
  • Interdependence of objects and their attributes: independent elements can never constitute a system
  • Goal seeking: systemic interaction must result in some goal or final state
  • Inputs and outputs: in a closed system inputs are determined once and constant; in an open system additional inputs are admitted from the environment
  • Transformation of inputs into outputs: the process by which the goals are obtained
  • Entropy: the amount of disorder or randomness present in any system
  • Regulation: a method of feedback is necessary for the system to operate predictably
  • Hierarchy: complex wholes are made up of smaller subsystems
  • Differentiation: specialized units perform specialized functions
  • Equifinality: alternative ways of attaining the same objectives (convergence)
  • Multifinality: attaining alternative objectives from the same inputs (divergence)

Defining organisms as biological system with behavioral output

Evolution, the effects of time passing, the life histories of systems and their parts, evolutionary systems are dynamic and complex systems with the capacity to evolve over time, examine evolutionary systems requires an interdisciplinary perspectives.

Evolutionary organisms can be defined as biological systems of subsystems:

A species is made of multiple individual organisms genetically similar with compatible reproduction systems – male and female:

Species can be define as a group of individual organisms with compatible reproduction system that make up an interbreeding group and in most cases very similar subsystems, organisms are considered to be in the same species if they can breed procreate and produce fertile offspring.

A species continuation is centered on the organization of males and females in reproduction group:

  • Interspecies: If the interaction is with members of a different species (as in the case of defending from a predator)
  • Intraspecies: If the interaction is with members of the same species (as in the case fighting for the opportunity to breed).
Organs are the biological modules and biomechanics parts which makes the subsystems in an organism system

Organism internal system is made of a collective of organs working together in the execution of a specific body function (biological system or body system). The functions of organ systems often share significant overlap. For instance, the nervous and endocrine system both operate via a shared organ, the hypothalamus. For this reason, the two systems are combined and studied as the neuroendocrine system. The same is true for the musculoskeletal system because of the relationship between the muscular and skeletal systems.

Organ systems and modules:
  • Organs of the digestive system: organs and structures not part of the alimentary canal that aid in digestion; they include the teeth, salivary glands, liver, gallbladder, and pancreas.
  • Organ of Corti: the organ lying against the basilar membrane in the cochlear duct, containing special sensory receptors for hearing, and consisting of neuroepithelial hair cells and several types of supporting cells.
  • Effector organ: a muscle or gland that contracts or secretes, respectively, in direct response to nerve impulses.
  • Enamel organ: a process of epithelium forming a cap over a dental papilla and developing into the enamel.
  • End organ end-organ: a nerve ending in which the terminal nerve filaments are encapsulated.
  • Golgi tendon organ: any of the mechanoreceptors arranged in series with muscle in the tendons of mammalian muscles, being the receptor for stimuli responsible for the lengthening reaction.
  • Reproductive organs female: the ovaries, fallopian tubes, uterus, vagina, and vulva (external genitalia) of a female. The breasts are a secondary sex character, enclosing the mammary glands.
  • Reproductive organs male: the external genitalia, accessory glands that secrete special fluids, and the accompanying ducts.
  • Sensory organs: organ that receive stimuli that give rise to sensations, i.e. organs that translate certain forms of energy into nerve impulses that are perceived as special sensations.
  • Target organ: organ that is affected by a particular hormone.
  • Vestigial organ: an undeveloped organ that, in the embryo or in some remote ancestor, was well developed and functional.

 

The systems of life are made of two main components

There are two main groups of organisms, Autotrophic organism or autotroph and Heterotrophic organisms or heterotroph (summarized from wiki):

Autotroph or primary producer: is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) from simple substances present in its surroundings, generally using energy from light (photosynthesis) or inorganic chemical reactions (chemosynthesis).

Heterotroph (chemoorganoheterotrophs or organotrophs organism) or a consumer: is an organism that cannot manufacture its own food and instead obtains its food and energy by taking in organic substances, usually plant or animal matter. All animals, protozoans, fungi, and most bacteria are heterotrophs.

Autotrophs are fundamental to the food chains of all ecosystems in the world. They take energy from the environment in the form of sunlight or inorganic chemicals and use it to create energy-rich molecules such as carbohydrates. This mechanism is called primary production. Heterotrophs, take in autotrophs as food to carry out functions necessary for their life.

All animals are heterotrophs and they are depending on autotrophs, or primary producers, for the energy and raw materials they need. Heterotrophs obtain energy by breaking down organic molecules (carbohydrates, fats, and proteins) obtained in food. Carnivorous organisms rely on autotrophs indirectly, as the nutrients obtained from their heterotroph prey come from autotrophs they have consumed. Most heterotrophs utilize organic compounds both as a carbon source and an energy source. Heterotrophs function as consumers in food chains: they obtain organic carbon by eating autotrophs or other heterotrophs. They break down complex organic compounds (e.g., carbohydrates, fats, and proteins) produced by autotrophs into simpler compounds (e.g., carbohydrates into glucose, fats into fatty acids and glycerol, and proteins into amino acids). They release energy by oxidizing carbon and hydrogen atoms present in carbohydrates, lipids, and proteins to carbon dioxide and water, respectively.

Most ecosystems are supported by the autotrophic primary production of plants that capture photons initially released by the sun. The process of photosynthesis splits a water molecule (H2O), releasing oxygen (O2) into the atmosphere, and reducing carbon dioxide (CO2) to release the hydrogen atoms that fuel the metabolic process of primary production. Plants convert and store the energy of the photon into the chemical bonds of simple sugars during photosynthesis. These plant sugars are polymerized for storage as long-chain carbohydrates, including other sugars, starch, and cellulose; glucose is also used to make fats and proteins. When autotrophs are eaten by heterotrophs (i.e. consumers) the carbohydrates, fats, and proteins contained in them become energy sources for the heterotrophs. Proteins can be made using nitrates, sulfates, and phosphates in the soil.

Autotrophs and heterotrophs compos the community of living organism and all the system of life

The optimal carrying capacity is the maximum spreading capacity of a biological species in an environment or the maximum population size (mass) of the species that the environment can sustain indefinitely for as long as abiotic and other environmental factors sustain, given the food, habitat, water, and other energetic or habitation necessities maintained.

Organism and their interaction with planetary systems of abiotic factors

Organisms properties are determined by their genetic code which in term determine the way they operate (behave), relative to the way other organisms in their community (all species in shared habitat) are operating, relative to the properties of the abiotic or the non-biological peripheral system of elemental and environmental systems of abiotic conditions and factors. The interaction between organism and abiotic systems are the predominant factor which affects the type and distribution of living organism on the planet. Abiotic systems include the following factors:

  • Temperature: affects metabolism, regulation of body temperature
  • Water and rainfall: affects desiccation of organisms
  • Sunlight: affects photosynthesis and photoperiods of primary producers
  • Rocks and soil, pH, mineral composition, physical structure: affect the distribution and properties of primary produces and consumers
  • Altitude and longitude: affects the atmospheric conditions of oxygen, sunlight radiation and temperature
  • Wind – amplifies temperature effects: increased evaporation: affects plant morphology
Abiotic factors determine ecozones and biome types

Ecozone is a biogeographic division of the Earth’s land surface, based on distributional patterns of terrestrial organisms and it characterized by a specific biome or biocommunities (the mix of organisms that coexist in the defined geographic area), there are no sharp boundaries between ecozones, in which a series of biocommunities display a continuous gradient) as they grade into each other.

Evolutionary processes and trends are the outcome of environmental conditions:

The rules of evolution are of a process of a change and bio-diversification where life is gaining ground when water, soil and climate supports a continuous expansion of biomass of the primary producers (plants) and with it, for as long as the supporting conditions maintained the bio-diversifying of consumers species will continue and new species will emerge and will evolve to adapt for specific niches as they are being created in the process until the community of life reaches a stage of a climax community where species evolution is halted in the optimum ecosystem EROEI (Energy Return On Energy Invested) such conditions are presented in the ecozones of the equatorial broadleaf rainforests.

The opposite process is happening when such optimal and regular conditions reversed and are trending in the opposite direction and causing a recession in ecosystems and such as when the precipitation is decreases and the production of biomass by the primary producers decline and as the recession continues the biodiversity of the community of consumers will decline and evolutionary process will start in many species as they are trying to adapt for different conditions in their habitat as they emerge, expands and extinct. When the biodiversity and the biomass of the primary producers is in decline, the consumers will follow in an evolutionary race to the bottom competing on the dwindling resources and habitats and favoring opportunism and rudimentary survival strategies.

An ecosystem can be in three stages:
  • Succession (development) is the expansion of the primary producers distribution, energy production, biomass storage and consumable energetic products (leaves, fruits, flowers etc.) creating a positive EROEI environment for the consumers – More energy is manufactured and captured by the primary producers as they expand their distribution, rang and growth captured as biomass where the community of consumers can evolve to directly and indirectly consume the energy captured in the biomass and the system is in a state positive EROEI and bio-diversification.
  • Equilibrium is a state of optimal production by the primary producers and maximal consumption by the consumers – When optimal conditions are maintained for a long period of time as in the case of the equatorial rainforests, the primary producers reach the maximal capacity of production and storage where the laws of physics and biology are stretched to the maximum. The ecosystem is now in a state of climax community and energy is balanced and maintained within the community in the form of stable biomass of the primary producers with maximum production, distribution and regularity of their products.
  • Recession is a state of contraction of the primary producer’s distribution, energy production, biomass storage and consumable energetic products creating a negative EROEI environment for the consumers – Less energy is manufactured and captured by the primary producers then the current community of consumers’ needs to consume in order to maintain climax community structure.

Evolutionary changes take place over long period of time in multiple generations of sexual selection with accordance to environmental conditions and their trends, in order to track the evolution of species including our own lineage one should factor the environmental conditions and trends and their dynamic interactions with our lineage ancestor’s developmental stage properties in each period of time:

  • Environmental conditional systems: atmospheric/climatic conditions temp, humidity
  • Elemental properties systems: distribution and availability of elements (sun, soil, stone, water etc.)
  • Structural systems: landscape, altitude.
  • Habitat: system of biological and non-biological systems.
  • Ecosystem: all the communities and the abiotic features of the environment
  • Biomes and ecozones: major types of communities often named for the predominant vegetation but also characterized by animals adapted to that environment.
  • Biosphere: that portion of earth inhabited by life and represents the sum of all communities and ecosystems, a closed system and largely self-regulating – the biosphere is the global ecological system integrating all living beings and their relationships, including their interaction with the elements of the planet:
    • Lithosphere: the earth’s crust and the portion of the upper mantle that behaves elastically on time scales of thousands of years or greater. The outermost shell of a rocky planet, the crust, is defined on the basis of its chemistry and mineralogy.
    • Hydrosphere: the combined mass of water found on, under, and above the surface of a planet.
    • Atmosphere: the layer of gases, commonly known as air that surrounds the planet Earth and is retained by Earth’s gravity. The atmosphere protects life on Earth by absorbing ultraviolet solar radiation, warming the surface through heat retention (greenhouse effect), and reducing temperature extremes between day and night (the diurnal temperature variations).

Ecological pyramids of energy production, stocking and exchange

Ecological pyramid (also trophic/eltonian/energy pyramid, food web/pyramid etc.): is a graphical representation designed to show the biomass or bio productivity at each trophic level in a given ecosystem.

Biomass: is the amount of stored energy in an ecological system in the living organisms that constructing it, biomass is the total amount of all living or organic matter presented in organisms in each level of the trophic pyramid and the total of all organisms that inhabit a defined area and entire regions of the same ecosystems/ecozone. Biomass pyramids show how much biomass is present in the organisms at each trophic level, while productivity pyramids show the production or turnover in biomass.

Energy pyramids begin with producers on the bottom (such as plants) and proceed through the various trophic levels (such as herbivores that eat plants, then carnivores that eat herbivores, then carnivores that eat those carnivores, and so on). The organisms that hold the highest level of the energetic hierarchies are considered to be at the top of the food chain (which is a term that has no objective value in the holistic view of things).

An energy pyramid of biomass shows the relationship between biomass and trophic level by quantifying the biomass present at each trophic level of an energy community at a particular time. It is a graphical representation of biomass (total amount of living or organic matter in an ecosystem) present in unit area in different tropic levels. Typical units are grams per square meter, or calories per square meter. One problem with biomass pyramids is that they can make a trophic level appear to contain more energy than it actually does. For example, all birds have beaks and skeletons, which despite having mass are not eaten by the next trophic level.

There is also pyramid of numbers which represent the number of organisms in each trophic level. They may be upright (e.g. Grassland ecosystem), inverted (parasitic ecosystem) or dumbbell shaped (forest ecosystem).

Pyramid of productivity and methods for measuring and presenting ecosystems states and properties

An ecological pyramid of productivity’ is often more useful, showing the production or turnover of biomass at each trophic level, instead of showing a single snapshot in time, productivity pyramids show the flow of energy through the food chain. Typical units are grams per square meter per year or calories per square meter per year.

As with the other ecological pyramids, a presentation of the pyramid of productivity in a graph will show the producers at the bottom and higher trophic levels on top.

When an ecosystem is healthy, this graph produces a standard ecological pyramid. This is because in order for the ecosystem to sustain itself, there must be more energy at lower trophic levels than there is at higher trophic levels. This allows organisms on the lower levels to not only to maintain a stable population, but also to transfer energy up the pyramid. The exception to this generalization is when portions of a food web are supported by inputs of resources from outside the local community. In small, forested streams, for example, the volume of higher levels is greater than could be supported by the local primary production. When energy is transferred to the next trophic level, typically only 10% of it is used to build new biomass and becoming stored energy (the rest going to metabolic processes). In this case, in the pyramid of productivity each step will be 10% the size of the previous step (…100,000, 10,000, 1,000, 100, 10, 1, 0.1, 0.01…).

The advantages of the pyramid of productivity as a representation:
  • It takes into account of the rate of production over a period of time.
  • Two species of comparable biomass may have very different life spans. Thus a direct comparison of their total biomasses is misleading, but their productivity is directly comparable.
  • The relative energy chain within an ecosystem can be compared using pyramids of energy; also different ecosystems can be compared.
  • There are no inverted pyramids.
  • The input of solar energy can be added.
The disadvantages of the pyramid of productivity as a representation:
  • The rate of biomass production of an organism is required, which involves measuring growth and reproduction through time.
  • There is still the difficulty of assigning the organisms to a specific trophic level. As well as the organisms in the food chains there is the problem of assigning the decomposers and detritivores to a particular trophic level.
  • Nonetheless, productivity pyramids usually provide more insight into an ecological community when the necessary information is available.
The three basic ways in which organisms get food are as producers, consumers and decomposers.

Producers (autotrophs) are typically plants or algae. Plants and algae do not usually eat other organisms, but pull nutrients from the soil or the ocean and manufacture their own food using photosynthesis. For this reason, they are called primary producers. In this way, it is energy from the sun that usually powers the base of the food chain. An exception occurs in deep-sea hydrothermal ecosystems, where there is no sunlight. Here primary producers manufacture food through a process called chemosynthesis.

Consumers (heterotrophs) are species that cannot manufacture their own food and need to consume other organisms. Animals that eat primary producers (like plants) are called herbivores. Animals that eat other animals are called carnivores, and animals that eat both plant and other animals are called omnivores.

Decomposers (detritivores) break down dead plant and animal material and wastes and release it again as energy and nutrients into the ecosystem for recycling. Decomposers, such as bacteria and fungi (mushrooms), feed on waste and dead matter, converting it into inorganic chemicals that can be recycled as mineral nutrients for plants to use again.

In real world ecosystems, there is more than one food chain for most organisms, since most organisms eat more than one kind of food or are eaten by more than one type of predator. A diagram that sets out the intricate network of intersecting and overlapping food chains for an ecosystem is called its food web. Decomposers are often left off food webs, but if included, they mark the end of a food chain. Thus food chains start with primary producers and end with decay and decomposers. Since decomposers recycle nutrients, leaving them so they can be reused by primary producers, they are sometimes regarded as occupying their own trophic level.

 

States, trends and properties of ecosystem

Equilibrium (climax community)

Climax community, or climatic climax community, is a historic term that expressed a biological community of plants and animals and fungi which, through the process of ecological succession the development of vegetation in an area over time, had reached a steady state. This equilibrium occurs because the climax community is composed of species best adapted to average conditions in that area. A “steady state” is more apparent than real, particularly if long-enough periods of time are taken into consideration.

Non-equilibrium by an Intermediate (transitional) disturbance

In intermediate disturbance, local species diversity is maximized when ecological disturbance is neither too rare nor too frequent. At high levels of disturbance all species are at risk of going extinct. At intermediate levels of disturbance, diversity is thus maximized because species that thrive at both early and late successional stages can coexist.

Low intermediate disturbance leads to decreased diversity while high intermediate disturbance causes an increase in species movement.

Intermediate disturbance levels would be the optimal amount of disorder within an ecosystem. Once K-selected and r-selected species (more about K and r selected species below) can live in the same region, species richness can reach its maximum. The main difference between both types of species is their growth and reproduction rate. These characteristics attribute to the species that thrive in habitats with higher and lower amounts of disturbance. K-selected species generally demonstrate more competitive traits. Their primary investment of resources is directed towards growth, causing them to dominate stable ecosystems over a long period of time; an example of K-selected species the African elephant, which is prone to extinction because of their long generation times and low reproductive rates. In contrast, r-selected species colonize open areas quickly and can dominate landscapes that have been recently cleared by disturbance. Ideal examples of r-selected groups are algae. Based on the contradictory characteristics of both of these examples, areas of occasional disturbance allow both r and K species to benefit by residing in the same area.

A non-equilibrium model is used to describe the relationship between disturbance and species diversity:
  • Ecological disturbances – have major effects on species richness within the area of disturbance
  • Interspecific competition – results from one species driving a competitor to extinction and becoming dominant in the ecosystem
  • Moderate ecological scale disturbances – prevent interspecific competition
Disturbances disrupt stable ecosystems and clear species’ habitat

Disturbances in ecosystem lead to species movement into the newly cleared area. Once an area is cleared there is a progressive increase in species richness and competition takes place again. Once disturbance is removed, species richness decreases as competitive exclusion increases.

Succession that begins in new habitats, uninfluenced by pre-existing communities is called primary succession, whereas succession that follows disruption of a pre-existing community is called secondary succession.

The trajectory of successional change can be influenced by many factors such as the site conditions, the character of the events initiating succession (perturbations), the Interactions between the present species and stochastic factors such as availability of colonists or seeds or weather conditions at the time of disturbance. Some of these factors contribute to predictability of succession dynamics; others add more probabilistic elements. It is a phenomenon or process by which an ecological community undergoes more or less orderly and predictable changes following a disturbance or the initial colonization of a new habitat.

Succession may be initiated either by formation of new, unoccupied habitat, such as from a lava flow or a severe landslide, or by some form of disturbance of a community, such as from a fire, severe windthrow or flood.

Successional dynamics beginning with colonization of an area that has not been previously occupied by an ecological community, such as newly exposed rock or sand surfaces, lava flows, newly exposed glacial tills, etc., are referred to as primary succession. The stages of primary succession include pioneer plants (lichens and mosses), grassy stage, smaller shrubs, and trees. Animals begin to return when there is food there for them to eat. When it is a fully functioning ecosystem, it has reached the climax community stage.

Successional dynamics following severe disturbance or removal of a pre-existing community are called secondary succession. Dynamics in secondary succession are strongly influenced by pre-disturbance conditions, including soil development, seed banks, remaining organic matter, and residual living organisms. Because of residual fertility and pre-existing organisms, community change in early stages of secondary succession can be relatively rapid.

Forest succession

The forests, being an ecological system, are subject to the species succession process. There are “opportunistic” or “pioneer” species that produce great quantities of seed that are disseminated by the wind, and therefore can colonize big empty extensions. They are capable of germinating and growing in direct sunlight. Once they have produced a closed canopy, the lack of direct sun radiation at soil makes it difficult for their own seedlings to develop. It is then the opportunity for shade-tolerant species to become established under the protection of the pioneers. When the pioneers die, the shade-tolerant species replace them.

  • Because of their higher reproductive rates and ecological opportunism, primary colonizers typically are r-strategists and they are followed by a succession of increasingly competitive flora and fauna.
  • The ability of an environment to increase energetic content, through photosynthetic capture of solar energy, increases with the increase in complex biodiversity as r species proliferate to reach a peak possible with K strategies.
  • r-strategists gradually being replaced by K-strategists which are more competitive and better adapted to the emerging micro-environmental characteristics of the landscape.
  • Eventually a new equilibrium is approached (a climax community) and biodiversity is now considered maximized at this stage, with introductions of new species resulting in the replacement and local extinction of endemic species.
Succession types – primary, secondary and cyclic

Ecological succession is the observed process of change in the species structure of an ecological community over time. The time scale can be decades (for example, after a wildfire), or even millions of years after a mass extinction. The community begins with relatively few pioneering plants and animals and develops through increasing complexity until it becomes stable or self-perpetuating as a climax community. The ʺengineʺ of succession, the cause of ecosystem change, is the impact of established species upon their own environments. A consequence of living is the sometimes subtle and sometimes overt alteration of one’s own environment.

Secondary succession

Secondary succession includes responses to natural disturbances such as fire, flood, and severe winds, and to Seasonal and cyclic dynamics

Cyclic succession

Unlike secondary succession, these types of vegetation change are not dependent on disturbance but are periodic changes arising from fluctuating species interactions or recurring events.

Causes of plant succession

Autogenic succession can be brought by changes in the soil caused by the organisms there. These changes include accumulation of organic matter in litter or humic layer, alteration of soil nutrients, change in pH of soil by plants growing there. The structure of the plants themselves can also alter the community. For example, when larger species like trees mature, they produce shade on to the developing forest floor that tends to exclude light-requiring species. Shade-tolerant species will invade the area.

Allogenic succession

Allogenic succession is caused by external environmental influences of nonliving factors in the environment and not by the vegetation. For example, soil changes due to erosion, leaching or the deposition of silt and clays can alter the nutrient content and water relationships in the ecosystems. Animals also play an important role in allogenic changes as they are pollinators, seed dispersers and herbivores. They can also increase nutrient content of the soil in certain areas, or shift soil about (as termites, ants, and moles do) creating patches in the habitat. This may create regeneration sites that favor certain species.

Trends in ecosystem and community properties during succession

Species diversity almost necessarily increases during early succession as new species arrive, but may decline in later succession as competition eliminates opportunistic species and leads to dominance by locally superior competitors. Net Primary Productivity, biomass, and trophic properties all show variable patterns over succession, depending on the particular system and site.

Ecological pyramids of productivity and biomass: a snapshot in time of an ecological community.

Biomass pyramid shows the amount of biomass at each trophic level while productivity pyramid shows the production or turn-over in biomass at each trophic level.

The bottom of the pyramid represents the primary producers (autotrophs). The primary producers take energy from the environment in the form of sunlight or inorganic chemicals and use it to create energy-rich molecules such as carbohydrates. This mechanism is called primary production. The pyramid then proceeds through the various trophic levels to the apex predators at the top.

When energy is transferred from one trophic level to the next, typically only ten percent is used to build new biomass. The remaining ninety percent goes to metabolic processes or is dissipated as heat. This energy loss means that productivity pyramids are never inverted, and generally limits food chains to about six levels.

Terrestrial biomass generally decreases markedly at each higher trophic level (plants, herbivores, carnivores). Examples of terrestrial producers are grasses, trees and shrubs. These have a much higher biomass than the animals that consume them, such as deer, zebras and insects. The level with the least biomass is the highest predators in the food chain, such as foxes and eagles.

Plants are the primary producers at the bottom of the pyramid after them comes the primary consumers (herbivores) followed by the secondary consumers (predators) followed by tertiary consumers (carnivores) and so on, the biomass pyramid decreases markedly in number of species and individuals of a species at each higher level of the food chain pyramid.

Trophic properties – Trophic level of an organism is the position it occupies in a food chain

A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism is from the start of the chain is a measure of its trophic level.

  • Trophic level 1 primary producers: plants and algae
  • Trophic level 2 primary consumers: herbivores eat plants
  • Trophic level 3 secondary consumers: predators eat herbivores
  • Trophic level 4 tertiary consumers: carnivores eat other carnivores
  • Trophic level 5 the top of the food chain: apex predators that have no predators

The path along the chain can form either a one-way flow or a food “web”. Ecological communities with higher biodiversity form more complex trophic paths.

The process of primary production by heterotrophic organisms

Primary production is the production of chemical energy in organic compounds by living organisms. The main source of this energy is sunlight but a minute fraction of primary production is driven by lithotrophic organisms using the chemical energy of inorganic molecules.

Regardless of its source, this energy is used to synthesize complex organic molecules from simpler inorganic compounds such as carbon dioxide (CO2) and water (H2O).
In the end point is a polymer of reduced carbohydrate, (CH2O), typically molecules such as glucose or other sugars. These relatively simple molecules may be then used to further synthesize more complicated molecules, including proteins, complex carbohydrates, lipids, and nucleic acids, or be respired to perform work. Consumption of primary producers by heterotrophic organisms, such as animals, then transfers these organic molecules (and the energy stored within them) up the food web, fueling all of the Earth’s living systems.

When a plant grows, new organic matter is created by the process of photosynthesis, which converts light energy into energy stored in chemical bonds within plant tissue. This energy fuels the metabolic machinery of the plant. New compounds and structures are synthesized, cells divide, and the plant grows in size over time. A plant requires sunlight, carbon dioxide, water, and nutrients, and through photosynthesis the plant produces reduced carbon compounds and oxygen.

Primary production definition: the synthesis and storage of organic molecules during the growth and reproduction of photosynthetic organisms.

Water is “consumed” in plants by the processes of photosynthesis (see above) and transpiration. The latter process (which is responsible for about 90% of water use) is driven by the evaporation of water from the leaves of plants. Transpiration allows plants to transport water and mineral nutrients from the soil to growth regions, and also cools the plant. Diffusion of water vapors out of a leaf, the force that drives transpiration, is regulated by structures known as stomata. These structures also regulate the diffusion of carbon dioxide from the atmosphere into the leaf, such that decreasing water loss (by partially closing stomata) also decreases carbon dioxide gain. Certain plants use alternative forms of photosynthesis, called Crassulacean acid metabolism (CAM) and C4. These employ physiological and anatomical adaptations to increase water-use efficiency and allow increased primary production to take place under conditions that would normally limit carbon fixation by C3 plants (the majority of plant species).

To calculate primary production we can measure the rate at which photosynthesis occurs as well as the rate in which the individual plant increases in mass when new chemical compounds and new plant tissue are produced. Over time, primary production results in the addition of new plant biomass to the system. Consumers derive their energy from primary producers, either directly (herbivores, some detritivores), or indirectly (predators, other detritivores).

Primary production on land is a function of many factors

Primary production has an upper limit to the amount of energy that can be captured by autotrophs the limits arise from abiotic factors:
Only a small fraction of the sun’s radiation is actually used in the photosynthetic reaction in plants at the Earth’s surface. Of the total solar radiation striking the Earth’s outer atmosphere, about half of it is reflected back to space by ice, snow, oceans, or deserts, or absorbed by gases in the atmosphere such as the ozone gas layer which absorbs nearly all ultraviolet light, which makes up about 9% of the sun’s radiation.
The autotrophs primary production limit is determine by multiple factors such as the intensity of solar radiation reaching the earth’s surface which depends geographical location defining the angle of the surface to the sun (the maximum energy intensity is received at the equator, and the intensity decreases as we move toward the poles), the altitude (higher intensity in higher altitude and vice-a-versa), atmospheric conditions such as clouds and the amount of humidity and particle in the atmosphere, daily exposure time to radiation (a mountain ridge can shade an area for few hours a day), there are also limits to primary production of biomass that arise due to other factors which involved in the process of photosynthesis such as the local hydrology the availability of water as well temperature, minerals in the ground and many more factors limit the primary production in geographic areas, and these differences have profound effects on observed geographic patterns of biomes in macro and micro climate zones.

Overall, of the light that reaches Earth’s surface, only about half of it is in the wavelength range that can be used by plants in photosynthesis (~400-700nm wavelength) – this is called the Photosynthetically Active Radiation (PAR). Plants strongly absorb light of blue and red wavelengths (hence their green color, the result of reflection of green wavelengths), as well as light in the far infrared region, and they reflect light in the near infrared region. Even if the wavelength is correct, the light energy is not all converted into carbon by photosynthesis. Some of the light misses the leaf chloroplast, where the photosynthetic reactions occur, and much of the energy from light that is converted by photosynthesis to carbon compounds is used up in keeping the plant biochemical “machinery” operating properly – this loss is termed “respiration”, although it also includes thermodynamic losses.

Plants do not use all of the light energy theoretically available to them, on average, plant gross primary production on earth is about 5.83 x 106 cal m-2 yr-1. This is about 0.06% of the amount of solar energy falling per square meter on the outer edge of the earth’s atmosphere per year (defined as the solar constant and equal to 1.05 x 1010 cal m-2 yr-1). After the costs of respiration, plant net primary production is reduced to 4.95 x 106 cal m-2 yr-1, or about 0.05% of the solar constant: this is the “average” efficiency, and in land plants this value can reach ~2-3% and in aquatic systems this value can reach ~1%. This relatively low efficiency of conversion of solar energy into energy in carbon compounds sets the overall amount of energy available to heterotrophs at all other trophic levels.

Gross primary production and net primary production

Gross primary production (GPP) is the amount of chemical energy as biomass that primary producers create in a given length of time. (GPP is sometimes confused with gross primary productivity, which is the rate at which photosynthesis or chemosynthesis occurs.) Some fraction of this fixed energy is used by primary producers for cellular respiration and maintenance of existing tissues (i.e., “growth respiration” and “maintenance respiration”).

The remaining fixed energy (i.e. mass of photosynthate in the form of sugar or other substance made by photosynthesis) is referred to as net primary production (NPP). Net primary production is the rate at which all the plants in an ecosystem produce net useful chemical energy; it is equal to the difference between the rate at which the plants in an ecosystem produce useful chemical energy (GPP) and the rate at which they use some of that energy during respiration. Some net primary production goes toward growth and reproduction of primary producers, while some is consumed by herbivores.

Both gross and net primary production are in units of mass per unit area per unit time interval.

In terrestrial ecosystems, mass of carbon per unit area per year is most often used as the unit of measurement.

Definitions and calculation methods of primary production

Gross Primary Production (GPP): is the total amount of CO2 that is fixed by the plant in photosynthesis. Respiration (R): is the amount of CO2 that is lost from an organism or system from metabolic activity. Respiration can be further divided into components that reflect the source of the CO2:

  • Rp : Respiration by Plants (Autotrophs)
  • Rh : Respiration by Heterotrophs
  • Rd : Respiration by Decomposers (the microbes)
Net Primary Production (NPP):

NEP is the net amount of primary production after the costs of plant respiration are included. Net Primary Production (NPP) = the Gross Primary Production (GPP) minus Respiration (R) or NPP = GPP – R

Net Ecosystem Production (NEP):

NEP is the net amount of primary production after the costs of respiration by plants, heterotrophs, and decomposers are all included. Net Ecosystem Production = the Gross Primary Production (GPP) minus the sum of all types of Respiration (Rp + Rh + Rd) or NEP = GPP – (Rp + Rh + Rd)

These definitions are only for “primary” and not “secondary” production

Secondary production is the gain in biomass or reproduction of heterotrophs and decomposers. The rates of secondary production are very much lower than the rates of primary production.

Measuring Primary Production is conduct by two general approaches:
  • The first one is measuring the rate of photosynthesis
  • The second one is measuring the rate of increase in plant biomass
Rate of Photosynthesis:

The process of photosynthesis utilizes carbon dioxide + water reaction with sunlight to produce glucose + oxygen + water. A balanced chemical equation for the process can be written as 6CO2 + 6H2O —> C6H12O6 + 6O2. Overall, photosynthesis uses light energy to convert carbon dioxide into a carbohydrate.

Theoretically we can measure the depletion of CO2 from plants tissue into the surroundings per unit time (or the generation of O2) to calculate the primary production. When plants exposed to light photosynthesis or primary production occur and in the absence of light photosynthesis or primary production do not accrue in both cases respiration maintained.

Cellular respiration occur throughout the daily cycle (light and darkness) and it is the reverse process from photosynthesis, a balanced chemical equation for the process can be written as C6H12O2 —> 6CO2 + 6H2O

Photosynthesis stores energy and respiration releases it for use in functions such as reproduction and basic maintenance. When calculating the amount of energy that a plant stores as biomass, which is then available to heterotrophs, we must subtract plant respiration costs from the total primary production.

Biomass – the mass of living biological organisms in a given area or ecosystem at a given time.

Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms, plants or animals. The mass can be expressed as the average mass per unit area, or as the total mass in the community.

How biomass is measured depends on why it is being measured. Sometimes, the biomass is regarded as the natural mass of organisms in situ, just as they are. For example, in a salmon fishery, the salmon biomass might be regarded as the total wet weight the salmon would have if they were taken out of the water. In other contexts, biomass can be measured in terms of the dried organic mass, so perhaps only 30% of the actual weight might count, the rest being water. For other purposes, only biological tissues count, and teeth, bones and shells are excluded. In stricter scientific applications, biomass is measured as the mass of organically bound carbon (C) that is present.

Apart from bacteria, the total live biomass on Earth is about 560 billion tones C, and the total annual primary production of biomass is just over 100 billion tones C/yr. However, the total live biomass of bacteria may exceed that of plants and animals. The total amount of DNA base pairs on Earth, as a possible approximation of global biodiversity, is estimated at 5.0 x 1037, and weighs 50 billion tones. In comparison, the total mass of the biosphere has been estimated to be as much as 4 TtC (trillion tons of carbon).

Different methods used for calculating the rate of biomass accumulation:

Calculating biomass dry weight (without the water) of plant material (stems, leaves, roots, flowers and fruits, minus the mass of the seeds that may have blown away) that was accumulated over one year in a defined area is one of the common methods for measuring production of biomass in terrestrial fauna.
The measure of primary production by rate of biomass accumulation is calculated in grams m-2 yr-1

There are specific methods for calculating biomass of certain types of ecosystems such as:
Grasslands biomass calculation methods

Most frequently, peak standing biomass is assumed to measure NPP. In systems with persistent standing litter, live biomass is commonly reported. Measures of peak biomass are more reliable if the system is predominantly annuals. However, perennial measurements could be reliable if there were a synchronous phenology (regularly recurring biological phenomena) driven by a strong seasonal climate. These methods may underestimate ANPP in grasslands by as much as 2 (temperate) to 4 (tropical) fold. Repeated measures of standing live and dead biomass provide more accurate estimates of all grasslands, particularly those with large turnover, rapid decomposition, and interspecific variation in timing of peak biomass. Wetland productivity (marshes and fens) is similarly measured.

Forests biomass calculation methods

Methods used to measure forest productivity are more diverse than those of grasslands. Biomass increment based on stand specific allometry plus litterfall is considered a suitable although incomplete accounting of above-ground net primary production (ANPP). Field measurements used as a proxy for ANPP include annual litterfall, diameter or basal area increment (DBH or BAI), and volume increment.

 

Animals

All animals are heterotrophs: they must ingest other organisms or their products for sustenance. Animals are multicellular, eukaryotic organisms of the kingdom Animalia (also called Metazoa). The animal kingdom emerged as a clade within Apoikozoa as the sister group to the choanoflagellates. Most known animal phyla appeared in the fossil record as marine species during the Cambrian explosion, about 542 million years ago. Animals can be divided broadly into vertebrates and invertebrates:

  • Vertebrates have a backbone or spine (vertebral column): They include fish, amphibians, reptiles, birds and mammals and amount to less than five percent of all described animal species.
  • Invertebrates lack a backbone: They include molluscs (clams, oysters, octopuses, squid, snails); arthropods (millipedes, centipedes, insects, spiders, scorpions, crabs, lobsters, shrimp); annelids (earthworms, leeches), nematodes (filarial worms, hookworms), flatworms (tapeworms, liver flukes), cnidarians (jellyfish, sea anemones, corals), ctenophores (comb jellies), and sponges.

The state of an animal species and its evolutionary trend are relative to the states and trends of its ecological community and to the species genotype

There are two fundamental states for a species, for individual organisms and for ecological community’s state of equilibrium and state of non-equilibrium.

The ecosystem’s non-equilibrium states have two options recession or development; accordingly a species or an individual organism can either have as a result a negative or positive short-term or long-term EROEI.

The state of an organism and a species is determine by its energetic stock that is available for output (work) and that is needed to be maintained by a specific energetic input, the dynamic interactions between the abiotic factors and individual organisms and multi-organism systems of energetic stocks are the defining factors of an ecosystem.

The evolution of motile animals

Motility

Motility is the function of an organism to move spontaneously and independently from one point in space to another point in space.
In biology, motility is the ability to move spontaneously and actively for a biological purpose (e.g. for consuming objects of energy (feeding), for copulating (reproduction), to move away (escape) from threats and hazards etc.). Motility is genetically determined and is determine by genotypic assets of motile organs and it is determined by the historic process of adaptation for environmental factors. The term motile applies to unicellular, simple multicellular and complex organisms (animals), as well as to some mechanisms of fluid flow in multicellular organs, in addition to animal locomotion.

Motility evolved in few different stages based on evolving strategies that drove motility for certain behavioral function first based on the fundamental existential domains for living organisms:

  • Maintaining internal structure: to stay alive in the immediate term by avoiding threats of other organisms and environmental hazards.
  • Maintaining energetic balance: to stay alive for the intermediate term by consuming energy via predation of other organisms
  • Maintaining the species: to stay alive in the long term by passing the genetic data via sexual reproduction

The basic 3 domains of our early ancestors only got more complex as the accumulation of genetic data offered more fine-tuned systems for motile operations.

Motility is the result of the development of logic functions of motile behavioral outputs

Definition of function: A function is a mathematical relationship in which the values of a single dependent variable (or a system of variables) are determined by the values of one or more independent variables (or systems of variables). Function means the dependent variable is determined by the independent variable(s).

Motility is a system of functions of a non-equilibrium system of biological matter

Biological systems are physiological systems of living organisms and are made by the organization and manipulation of physical systems of chemical, electrical, mechanical of properties utilizing the governing physical laws. Physical and non-biological systems have only two states:

  • Equilibrium: a state of stability when EROEI is in a constant state of balance
  • Non-equilibrium: a state of instability when EROEI is in a state of equalization from a state of energetic surplus or energetic deficit.
Equilibrium

In thermodynamics equilibrium is relevant for non-living systems; it is an axiomatic concept of thermodynamics of an internal state of a single thermodynamic system (e.g. structure of uniform atoms, molecules etc.), or a relation between several thermodynamic systems connected by more or less permeable or impermeable walls (i.e. organization of different compounds of atoms and molecules crating systems of balanced thermal mechanical, chemical, or radiative exchange).
In thermodynamic equilibrium there are no net macroscopic flows of matter or of energy, either within a system or between systems. In a system in its own state of internal thermodynamic equilibrium, no macroscopic change occurs. Systems in mutual thermodynamic equilibrium are simultaneously in mutual thermal, mechanical, chemical, and radiative equilibria. Systems can be in one kind of mutual equilibrium, though not in others. In thermodynamic equilibrium, all kinds of equilibrium hold at once and indefinitely, until disturbed by a thermodynamic operation. In a macroscopic equilibrium, almost or perfectly exactly balanced microscopic exchanges occur; this is the physical explanation of the notion of macroscopic equilibrium.

A thermodynamic system in its own state of internal thermodynamic equilibrium has a spatially uniform temperature. Its intensive properties, other than temperature, may be driven to spatial inhomogeneity by an unchanging long range force field imposed on it by its surroundings.

In non-equilibrium systems, by contrast, there are net flows of matter or energy. If such changes can be triggered to occur in a system in which they are not already occurring, it is said to be in a metastable equilibrium.

Though it is not a widely named law, it is an axiom of thermodynamics that there exist states of thermodynamic equilibrium. The second law of thermodynamics states that when a body of material starts from an equilibrium state, in which portions of it are held at different states by more or less permeable or impermeable partitions, and a thermodynamic operation removes or makes the partitions more permeable and it is isolated, then it spontaneously reaches its own new state of internal thermodynamic equilibrium, and this is accompanied by an increase in the sum of the entropies of the portions.

One fundamental difference between equilibrium thermodynamics and non-equilibrium thermodynamics lies in the behavior of inhomogeneous systems, which require for their study knowledge of rates of reaction which are not considered in equilibrium thermodynamics of homogeneous systems.

Non-equilibrium systems

Non-equilibrium thermodynamic systems are physical systems that are not in thermodynamic equilibrium but can be described in terms of variables (non-equilibrium state variables) that represent an extrapolation of the variables used to specify the system in thermodynamic equilibrium. Non-equilibrium thermodynamics is concerned with transport processes and with the rates of chemical reactions. It relies on what may be thought of as more or less nearness to thermodynamic equilibrium.

Difference between equilibrium and non-equilibrium thermodynamics

A profound difference separates equilibrium from non-equilibrium thermodynamics. Equilibrium thermodynamics ignores the time-courses of physical processes. In contrast, non-equilibrium thermodynamics attempts to describe their time-courses in continuous detail.

Equilibrium thermodynamics restricts its considerations to processes that have initial and final states of thermodynamic equilibrium; the time-courses of processes are deliberately ignored. Consequently, equilibrium thermodynamics allows processes that pass through states far from thermodynamic equilibrium that cannot be described even by the variables admitted for non-equilibrium thermodynamics such as time rates of change of temperature and pressure. For example, in equilibrium thermodynamics, a process is allowed to include even a violent explosion that cannot be described by non-equilibrium thermodynamics. Equilibrium thermodynamics does, however, for theoretical development, use the idealized concept of the “quasi-static process”. A quasi-static process is a conceptual (timeless and physically impossible) smooth mathematical passage along a continuous path of states of thermodynamic equilibrium. It is an exercise in differential geometry rather than a process that could occur in actuality.

Non-equilibrium state variables

The suitable relationship that defines non-equilibrium thermodynamic state variables is as follows:

When a system happens to be in states that is sufficiently close to thermodynamic equilibrium, non-equilibrium state variables are such that they can be measured locally with sufficient accuracy by the same techniques as are used to measure thermodynamic state variables, or by corresponding time and space derivatives, including fluxes of matter and energy.

In general, non-equilibrium thermodynamic systems are spatially and temporally non-uniform, but their non-uniformity still has a sufficient degree of smoothness to support the existence of suitable time and space derivatives of non-equilibrium state variables. Because of the spatial non-uniformity, non-equilibrium state variables that correspond to extensive thermodynamic state variables have to be defined as spatial densities of the corresponding extensive equilibrium state variables. On occasions when the system is sufficiently close to thermodynamic equilibrium, intensive non-equilibrium state variables, for example temperature and pressure, correspond closely with equilibrium state variables. It is necessary that measuring probes be small enough, and rapidly enough responding, to capture relevant non-uniformity. Further, the non-equilibrium state variables are required to be mathematically functionally related to one another in ways that suitably resemble corresponding relations between equilibrium thermodynamic state variables.

Living organisms and communities of living organisms are non-equilibrium systems

All biological systems by definition are not in thermodynamic equilibrium; for they are changing or can be triggered to change over time, and are continuously and discontinuously subject to flux of matter and energy to and from other systems and to chemical reactions.

Defining a conceptual framework for a non-equilibrium biological systems based on thermodynamics principles used for modeling non-biological systems

Non-equilibrium thermodynamics is attempting to describe continuous time-courses, need its state variables to have a very close connection with those of equilibrium thermodynamics. Utilizing non-equilibrium thermodynamics principals for dealing with biological and evolutionary concepts requires the development of conceptual framework.

Metastability – the operator of the behavioral function of biological systems

In physics, metastability denotes the phenomenon when a dynamical system spends an extended time in a configuration other than the system’s state of least energy. During a metastable state of finite lifetime, all state-describing parameters reach and hold a stationary value:

The state of least energy is the only one the system will inhabit for an indefinite length of time, until more external energy is added to the system (unique “absolutely stable” state);

The system will spontaneously leave any other state (of higher energy) to eventually return (after a sequence of transitions) to the least energetic state.

Isomerization is another strategy for achieving metastability. Higher energy isomers are long lived as they are prevented from rearranging to their preferred ground state by (possibly large) barriers in the potential energy.

The metastability concept originated in the physics of first-order phase transitions. It then acquired new meaning in the study of aggregated subatomic particles (in atomic nuclei or in atoms) or in molecules, macromolecules or clusters of atoms and molecules. Later, it was borrowed for the study of decision-making and information transmission systems.

Metastability is common in physics and chemistry – from an atom (many-body assembly) to statistical ensembles of molecules (viscous fluids, amorphous solids, liquid crystals, minerals, etc.) at molecular levels or as a whole (see metastable states of matter and grain piles below). The abundance of states is more prevalent as the systems grow larger and/or if the forces of their mutual interaction are spatially less uniform or more diverse.

In dynamic systems (with feedbacks from variables) like logic decisional systems and neuroscience – the time invariance of the active or reactive patterns with respect to the external influences defines stability and metastability, In these systems, the equivalent of thermal fluctuations in molecular systems is the “white noise” that affects signal propagation and the decision-making.

Logic is the mechanism of tactics to drive a non-equilibrium system into a state of metastability

Equilibrium state variables are the ones that measure the state of the internal systems (i.e. visceral perception) relative to the optimal state of a perceived equilibrium (the notion of the state) it is done by sensors that measure the level of entropy in the system (a quantity representing the unavailability of a system’s thermal energy for conversion into mechanical work, often interpreted as the degree of disorder or randomness in the system).

The somatosensory system is the system that was responsible for the assessment of the metastability in early organisms utilizing afferent sensory neurons and pathways that respond to changes at the surface or inside the body. The axons of sensory neurons connect with, or respond to, various receptor cells. These sensory receptor cells are activated by different stimuli giving a functional meme to the responding sensory neuron, such as a thermoreceptor which carries information about temperature changes. Other types include mechanoreceptors, chemoreceptors, and nociceptors and they send signals along a sensory nerve to the spinal cord where they may be processed by other sensory neurons and then relayed to the brain for further processing. Sensory receptors are found all over the body including the skin, epithelial tissues, muscles, bones and joints, internal organs, and the cardiovascular system.

Somatic senses are sometimes referred to as somesthetic senses, with the understanding that somesthesis includes the sense of touch, proprioception (sense of position and movement), and (depending on usage) haptic perception.

The mapping of the body surfaces in the brain is called a cortical homunculus and plays a fundamental role in the creation of body image. This brain-surface (i.e. cortical) map is not immutable, however. Dramatic shifts can occur in response to stroke or injury.

The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons (including the sensory receptor cells), neural pathways, and parts of the brain involved in sensory perception. Commonly recognized sensory systems are those for vision, hearing, touch, taste, smell, and balance. In short, senses are transducers from the physical world to the realm of the mind where we interpret the information, creating our perception of the world surrounding organisms.

The receptive field is the area of the body or environment to which a receptor organ and receptor cells respond. For instance, the part of the environment that an eye can intercept is its receptive field; the light that each rod or cone can see, is its receptive field. Receptive fields have been identified for the visual system, auditory system and somatosensory system.

The relative level of entropy is collected via the somatosensory system and other senses responsible for physical proprioception input creating a function declaration (defines a named function variable without requiring variable assignment) that can fit into one or more visceral discipline functions.

The output of the metastability state is a product of the physical proprioception systems that motivate the haptic perception for motile function:

The somatosensory is the entity responsible for the output of the statement in the form of haptic notion of the logic function, which can be negative or positive, if the notion is positive then the entropy state is closer to a state of equilibrium and no action is needed but if the notion is negative and the entropy levels are declining than an action is needed.

The notion is the signal for a motion which and the function is an action to regain metastability by executing a strategy of behaviors. The notion is the apparatus for the strategy and it is the operator of the haptic perception which is the

The sense of spatial orientation the first visceral function of motility

The sense of spatial orientation was the first notion setting variable for the first motile function of behavioral motility utilizing spatial orientation of isomers for the function of navigation in two opposite direction on a single dimension vertical axis – up or down.

The initial sense of spatial orientation was based on monitoring chemical reactions in hydrocarbons that create

The plain field for the movement was

The mechanism of motility in animals include a function of logic

Motility in animals is based on a linear mechanism of 3 functions (unlike the motile mechanism in visceral function that operated the motility in organism that was based on two functions the first is metastability declaration function and the second is execution of motile function which can be referred to as reflexive function):

Metastability state declaration function

Establish visceral perception declaration utilizing Somatic senses to measure concurrent internal state variables of an organism relative to a database of optimal state variables

The internal state variables are in two groups:

  • Measure of the internal physiological state of entropy
  • Measure of the external environmental variables contributing to the internal state of entropy
Planning of motile function

Identification of environmental variable via perceptual sensors that belongs to one of the three groups:

  • Variables that are causing or can cause a reduction of metastability (hazards, predators etc.)
  • Variables that can support an increase of metastability (nutrition)
  • Variable of motile mediums and environmental properties that can be used for execution of motile function
Execution of motile function

Establish haptic declaration of orientation and relativity to the environmental mediums and properties all the way to the nearest target of motile function (i.e. plan the rout from point A to Point B)

Operation of motile organs to perform the function of motility utilizing environmental medium and properties

Fundamental adaptations for fundamentally different environments – from living in the Hydrosphere to living on the Lithosphere and being dependent on the Atmosphere

A motile creature with adaptation to aquatic biosphere experience all the information in 3D and its life are limited to one medium – water: in this environment weightless floating and throttling fins are sufficient for movement and electrical, olfactory and visual sensors together with basic brain modules and basic capabilities are sufficient for fulfilling the tasks of orientation and navigation. In the aquatic environment fitness strategies do not rely on many brain parts and neurological circuits and all of the energetic needs are easily answered via rudimentary cognition.

Sharks for example maximized their evolutionary process with the design of the white shark et Al type of physical properties, efficient propulsion based locomotion and sensory and weaponry arsenal that can maintain surplus or equalized energetic balance and EROEI and thus the species reached the peak of it evolutionary process and the developmental diversifying into sister groups with different variations of later designs for niche adaptation are not affecting the prototype evolutionary stagnation.

The transition of an aquatic creature like the shark into a terrestrial animal like the wolf requires a fundamental adaptation to take place and a new fitness strategy must emerge and evolve in order to adjust to a fundamentally different environment of an fundamentally different properties of the abiotic mediums of air and being limited (at least in the beginning) to a habitat that include only 2 dimensional movement (left/right, forward/backward and all the angles in between) on surfaces of land and structures, new challenges arisen from the properties of the new mediums that construct the terrestrial environment:

Surrounded by Atmosphere (air/gas) instead of being submerged in the Hydrosphere (liquid): as the only medium for respiration, for sensory interception of photons with wider spectrum of wave frequencies, inferior acoustics and pressure wave transmit over distances with faster echoing and surfaces distortion, poor and chaotic carrier for high concentration wad of olfactory molecules but also with series hazards such as wider and stronger solar radiation with less protection from the hazardous infrared radiation and higher levels of ultraviolet radiation higher when climbing in altitudes, higher temperatures variability and extremes such as the temperature range of the water which its lower limit is above zero Celsius at sea level pressure vs. the lower range of terrestrial environments temperatures at sea level of −89.2 °C (−128.6 °F, 184.0 K), in other words aquatic animal do not have to deal with the risk of freezing because if that happened their environment would also be nulled, hydration presents a fundamental limitation in every terrestrial habitat. Other hazardous factors of the atmospheric environment arise by the flow and movement of the gas molecules between higher and lower barometric pressure areas such as the long range and wide spread of bacteria’s and other pathogens over geographic areas and many more abiotic hazards such as weather conditions, precipitation and others.

Living on the Lithosphere with the constant energetic burden of Gravity instead of energetically neutral weightless floatation: dependency on 2 dimensional surfaces aligned to gravity or gradable structures is presenting real challenges such as energetic challenges of movement and the cognitive challenge operating complex biomechanics and for orientation in the limiting environment and the challenges and limitations arise from the atmospheric medium being the only source for sensory information interception which is required for traveling on the surface. Living on the surface of objects in the lithosphere is presenting many hazardous threats and limitations when traveling: vertical structures that block the way and the view, vertical gaps where animals can fall into and all due to gravity.

A major adaptation is needed physically and the biomechanical parts are only one aspect of the adaptation, the second aspect is cognitive adaptation which must also profoundly evolve in order to operate such biomechanical functions under the energetic burden and limitations of gravity in an atmospheric environment which is especially crucial for a quadruped performing locomotion gaits and dynamically balancing the body (including other parts that not part of the biomechanics) to gravity in accordance to other thermodynamics forces which governing the movement as well as the arrangements and the sizes of the body parts (head, tail, digestive system etc.) which have to follow that fundamentals while taking in consideration for the design the energetic expenditure of thermoregulation and feeding strategy.

And although the physical challenges of changing fundamental environment of certain living medium and perceptual perspective are immense the bigger challenge is to tie all the biological development into operation and follow a fitness strategy which demands new cognitive strategy based on the new perspective for orienting and navigating in the 2 dimensional habitats.

Moving between fundamental environments is the main factor in species cognitive abilities the secondary one is the level of adaptation needed for reaching equilibrium in specific fundamental environment.

After a certain genotype of certain phenotypic design reached equilibrium in a certain environment it will spread to the extent of its environment boundaries the lineage will reach the maximum limitation of its ecosystem caring capacity, at this point a lineage evolution will stagnate but as conditions changes and new niches emerge it will start to develop into sister groups and branch into new variations with similar genotype and cognition but not necessarily similar phenotype of physical and behavioral characteristics (e.g. in aquatic environment sharks achieved fundamental adaptation and other type of more specialized sister species evolved such as nurse sharks, manta rays, hammer heads etc.).

A lineage reach the state of complete adaptation will be considered as outgroup and as new ecological niches emerge it will start branching into more specialized versions or sister group adapted for more specific nieces in the same fundamental environment.

Every time a lineage achieves a complete transition between two environment fundamentals the lineage also experienced a leap forward in its cognition.

The main factors in cognitive evolution of a species are the combination of the environment fundamentals and selected locomotion strategy for movement

The environmental fundamental adaptation factors:
  • The dimensions available movement in the environment – 2D vs. 3D
  • The complexity of the environment – the quantity, distribution and variation of objects and surfaces available for locomotion, orientation and navigation.
  • The distribution and location of energetically resources available for consumption.
The dimensions available for movement in an environment are the drivers for motile adaptation and the driver for cognitive evolution

The most fundamental drivers for movement in an environment are the number of available dimensions for movement, there are 2 types of fundamental environments:

  • 3 Dimension mediums – medium of bodies of water (e.g. oceans), medium of bodies gas (e.g. atmosphere) and medium of bodies of land mass (e.g. underground)
  • 2 Dimension environments – surfaces and structures (e.g. on top of ground, rocks, trees etc.)
The three dimensional (3D) environment movement fundamentals

Animals swimming in the water, flying in the air, digging tunnels under the ground surface or move between branches in a forest canapé have three dimensions available for their movement (X,Y and Z: up/down, forward/backward and left/right). All the animals with adaptation for 3D movement in 3D environment obeys the same fundamental rules

The two dimensional (2D) environment movement fundamentals

Animals moving on a surface (ground, branches, rocks, ocean bed etc.) have only two dimensions available for their movement (X and Y: forward/backward and left/ right). All of the other adaptations are just a derivative of these fundamental adaptations. All the animals adapted for 2D movement in 2D environment obeys the same fundamental rules

All organisms moving in 3 dimensions will obey the same environment movement fundamentals but their method of movement (locomotion) in their specific habitat may differ: fish and squid living in the 3D environment of water, bat and bird living in the 3D environment of gas (air) and monkey living in the 3D environment of the forest canapé will be subject to the same fundamentals of navigation and orientation in a 3D environment, while a dear, bear, snake and spider will obey the fundamental rules of navigation and orientation in a 2D environment.

Species primary adaptation strategy – the main fitness strategy to environmental fundamentals

A fundamental environment obtain a certain limitations that are dependent not only on its essential properties of the mediums of liquid or gas (air or water) which govern the crucial laws of physics and the rules thermodynamics for a movement in such environment but also by the structural complexity of such environment which determine the available methods for orientation, navigation and type of locomotion.

The different environmental fundamentals present different type of approaches and methods possibilities for a motile organism.

There are few factors in a species adaptation to a fundamental environment that will define the type of locomotion strategy such as the genetic assets and the availability, the distribution of the energetic resources and threats (which define the feeding strategy). The type of locomotion strategy can be based on agility – high speed movement or passivity – low speed movement and though will define the type and level of cognitive adaption.

The fundamental fitness strategy of a species is the methods of utilization of the environmental fundamental factors and that will determine its locomotion and its cognitive adaptation.

The basic rules of adaptation and transition between fundamental environments

Every time a species transit from one fundamental environment to another, the first main adaptation is a major cognitive change that is needed to achieve fitness when moving and navigating in certain environment: locomotion in a 3D environment and in a 2D environment demand very different cognitive tools.

The complexity or the simplicity of an environment structures and arrangement are drivers for cognitive evolution and “leap forward” in cognition

The main factors in cognitive leaps forward are the transition between fundamental environments:

  • Fish achieved rudimentary cognition for the only 3D environment they have evolved to live in
  • Birds and bats that evolved from pelagic animals to terrestrial animals and into volant animals (3D to 2D and to 3D) and have achieved a higher cognition due to the extreme limitation of the weight of their head and skeletons in order to fly (insect’s limited size is due to the breathing method of tracheae).
  • Land mammals (including mammals that walk on the surface of branches) that evolved from pelagic animals to terrestrial animals (3D to 2D) have achieved a higher cognition due to the moderate limitation of the weight of their head in order to be supported by their neck (the variation of their cognition is usually defined by the complexity of their environment)
  • Lesser apes such as Hylobatidae (gibbons) that use the trajectory brachiation locomotion evolved from pelagic animals to terrestrial animals and into type of arboreal-volant animal performing parabolic trajectories in high speed in the forest’s canapé (3D to 2D and to 3D) have achieved the fourth highest cognition due to their extreme agility and the complexity of their environment and due to the limitation of the weight of their head by such type of locomotion which can be achieved only if their total body weight is not too high for be supported by the branches (although hanging from branches align the head with the neck and the body to the center of gravity)
  • Ocean mammals like dolphins that evolved from pelagic animals to terrestrial animals and back into pelagic animals (3D to 2D and to 3D) have achieved the third highest cognition due to the lack of limitation of the weight of their head that is supported by the water but they didn’t evolve further due to the low complexity of their environment (sea lions and other animals that need to breed on land are still limited by the size and weight of the head)
  • Apes such as the pan members that use brachiation locomotion and quadruple locomotion evolved from pelagic animals to terrestrial animals to arboreal-volant animal and for semi-arboreal semi-terrestrial (3D to 2D to 3D to semi 2D i.e. living between the 3D of the canapé and the ground) achieved the second highest cognition due to the larger size of their brain relative to the larger size of their body and limited by their quadruped posture (which pressure their neck).
  • Humans that evolved from pelagic animals to terrestrial animals to arboreal-volant animal and back into terrestrial (3D to 2D to 3D and back to 2D) have achieved the highest non-rudimentary cognition due to the lack of limitation of the weight of their head as bipedal animal.

Achieving absolute adaptation to environment fundamentals (i.e. medium of existence, medium of movement and mediums of perception) creates lineages adaptation to certain locomotion design or certain feeding niche diversify the linage the definition of species and other classes of organisms is obscuring the dynamics of the process of evolution where linages of certain filament of DNA revolving in equilibrium or evolving and changing incrementally over generational of reproduction and pruning of unsuccessful or unlucky DNA sequence, the ones that were subdued to extinction by conditions changing unfavorable to their phenotypic expression.

The absolute-adaptation process is deemed as success (into certain locomotion design and perceptual medium of movement i.e. 3D/2D, Swimming/flying/brachiating/walking on four/two, movement medium of water/air/ground/underground etc.) if survival equilibrium was achieved by a certain sequence of DNA at this point the following process is the diversification of the DNA into more advanced forms capitalizing on the first successful design.

In any lineage there was one sequence of DNA which achieved the absolute-adaptation: for example there was one species in the human lineage that complete the geographical transit and physiological transition from the 3D environment of the oceans to the 2D environment of the land and by that complete absolute adaptation and reached equilibrium (a certain species of amphibian) from that point it started to branch into sister groups that obeys to the same fundamentals of the terrestrial 2D environment after that stage diversification emerge and certain strands of DNA mutate the designs of individuals and incrementally adapted different fitness strategies for moving around and performing feeding and reproducing in accordance to specific habitat’s niche’s attributes (e.g. resources, terrain etc.).

For a species, reaching adaptation for certain fundamental environment is the most challenging of all evolutionary processes as it demands also fundamental cognitive adaptation which is correlated to specific phenotypic properties and cognitive potential (i.e. the cognitive change in the simple nerve system of an insect becoming aquatic is not as major (due to genotypic limitations) as for large, long legged, otter-like mammal with big brain going the same path and becomes a whale).

If a lineage of animal never changed it environment after reaching the equilibrium, its brain will remain almost the same for as long as it is still lives in the same environment, same for all of its descendants and branches: they may have extreme size and weight variations but their cognition will be very similar until one branch will go through another environmental transition requiring fundamental adaptations, for example one fish species have reached equilibrium of design hundreds of million years ago and although since fish spread into almost every 3D underwater habitat niches, their cognition have stayed mostly unchanged since it reached the point of equilibrium in such fundamental environment, Dolphins on the other hand had the same evolutionary adaptation for 3D environment fundamentals as fish, but then they moved out of the ocean and adapt for living on the land in 2D environment, and once more moved to the oceans where again they adapt to 3D environment and although the phenotypic physics of Dolphins evolved to be very similar to fish, their genetic heritage of three transitions and adaptations can be observed in the differences between their brain size and complexity and their corresponding cognitions observed as phenotypic behaviors.

Fish have never changed their original fundamental environment and its cognition has stayed at the same since, the lineage of Dolphins currently classified as cetaceans, changed their fundamental environment 3 times and their cognition is highly developed as a result.

Dolphins’ intelligent is also supreme to all the land animals that went through 2 absolute-adaptations such as dogs and cats, and is in a similar level to animals reached the equilibrium point of the absolute-adaptation the same number of times – the Primates (water (3D), land (2D) and forest canapé (3D)), and all of their cognitions are inferior to the one cognition possessed by Humans which achieved 4 absolute-adaptations.

A species cognitive abilities and intelligence goes together with the number of transitions between fundamental environments as it accumulates more cognitive heritage in the process.

We are the species with the highest number of transitions between fundamental environments:

We had the same evolutionary adaptation for 3D environment in the early oceans, than we moved to the land and adapt for living in 2D environment, then we transit into the 3D environment of the forest canopy and again we landed on the ground and adapt for living in 2D environment.

New brain modules for new motile challenges presented by living on land

As our ancestors transition to land new cognitive system was developed to synchronize the mechanical parts and to direct the machine towards food and away from danger, with new or upgraded sensors to orient and to direct itself in the right direction and through the optimal path in order to locate food, mate or to distance itself away from enemies.

The new cognitive system was based on new and enhanced brain tissue and modules integrated into new or enhanced sensory modules creating new sensory based perceptual system.

The new cognitive system was very different and much more advanced than the previous aquatic cognitive system that was sufficient for maintaining equalized or positive energetic balance and EROEI in the 3 dimensional aquatic weightless floating in the pelagic zone easily throttling with minimal energetic expenditures. The liquid medium of motility activities, locomotion strategy and thermodynamics made it easy to travel efficiently and fast over large distances and landscapes in a bird view perspective, where the pelagic environment is always the same, with no need for thermoregulation due to the gradual and limited range of thermal fluctuations and steady temperatures in large zones of certain altitudes and depth a zone where all of the information for orientation and navigation and motile activities is easily intercepted directly from the highly conductive medium which compose their entire living habitat.

From wiki: “Pelagic fish live in the pelagic zone of ocean or lake waters – being neither close to the bottom nor near the shore – in contrast with demersal fish, which do live on or near the bottom, and reef fish, which are associated with coral reefs”

The transition from the fundamentals which are governing the phenotype of an animal in an aquatic environment, to the fundamentals governing the life of an animal in a terrestrial environment, demanding major morphological, physiological and cognitive adaptation.

When our ancestors transit into the land and developed traits for terrestrial environment and overcome the most important evolutionary obstacles of respiration, dehydration and thermoregulation, they also achieved an energetically and biomechanically efficient design of quadruple locomotion where the body is aligned vertically to gravity, supported by 4 limbs with the head and tail balancing each other horizontal spread of weight and opposing each other to oppose g-force force when changing course during gait.

Once the basic design was achieved evolution turned its focused on the haptic perception in the limitation of the 2 dimensional perspective where the censors are now in a medium that have no electrical conductivity, and the properties of gas reducing significantly the distances pressure waves (sound) can travel also the interception field available for capturing visual data and sound, are limited by the surrounding objects and surfaces.

The sensory perception of the pelagic animal went through stages in the process of transitioning from pelagic living to terrestrial living.

The first stage was transitioning from the pelagic lifestyle to demersal lifestyle; at this point the perceptual point of view evolved to answer the challenges arise from the limitations of the 2 dimension view field of an animal that living on vertical or horizontal surfaces on the bottom of the ocean, on ground or on the surfaces of objects (e.g. rocks, branches etc.).

The evolutionary stage of becoming demersal enable an easier evolutionary process of transition and adapting to terrestrial locomotion and orientation (without having the need yet for adapting to respiration of gas which came later), the demersal animal could gradually evolve for terrestrial locomotion while maintaining similar strategies for conducting motile activities as a surface animal on the seabed (together with the fact that there was no real threat to intervene in the fragile stage of adaptation locomotion in gravity when limited movement on the water edge makes the first pioneers an easy target for predation , and maybe that is the reason that it only happened at that time of the “D-day” of the animals invasion at the preliminary succession of the lithosphere.

Once the demersal animal completed the first requirement of respiration for terrestrial living the evolutionary focus shifted to more important adaptation of the locomotion in the unforgiving lithospheric environment ruled by a strong force of gravity demanding higher energetic expenditures for mobilizing increasing units of mass, and thermodynamics that govern increasing energetic expenditure for traveling over distances and for accelerating the speed of such mass and many other new hazards of the terrestrial environment such as finding water, falling from heights, and high range of thermal fluctuations daily and between weather conditions an seasons that can dehydrate or refrigerate all represented a new governing rules for energy sensitive systems where efficient locomotion with positive EROEI, thermoregulations and was a must for achieving fitness and for conducting all the existential activities.

In the apparatus of energy conservation strategy life started to adapt into amphibian animals and later for fully terrestrial animals which their adaptation process was driven by the vast availability of energy in the sugars produced by the photosynthesis of the early flora that was already developed for eons (and enriching the atmosphere with high levels of oxygen) and covered vast areas of the terrestrial land coupled with the nutrition of early colonizers such as insects that have already inhabit the early fauna.

The sensory perception that was suitable for living on the surface of water bodies needed also to adapt and ones the quadrupled design emerge as the optimal design for terrestrial locomotion other strategies emerge both physical and cognitive for enhancing its EROEI and functionality.

As the animals transition from water to land completed the focused first shifted to cognitive and sensory adaptation a new system for producing motile activities emerged and was dependent on new sensory systems based on smell, vision and sound with an increasing cognitive abilities based on new brain modules, produced more complex strategies for dealing with the lithospheric environment more demanding conditions.

The cognition was the main focus factor of the different fitness strategies of the early terrestrial animals rather than the morphology and the rudimentary control of the common phenotypic design which was shared in various variations by the new terrestrial linages of both our ancestors the early mammals and by the reptiles that inherit the world after them.

The variations that can be observed are insignificant changes in the external quadruple design such as bigger bodies, different proportions, postures and muscles, differences in the head and tail and other adaptations to different feeding strategies and habitat niches, other variations are in phenotypic expressions of claws, teeth such as stronger jaws or limbs with bigger and longer claws and fangs in carnivore for the purpose of predation, yet not fundamentally different in their functionality, design and cognitive management than the one that they are preying on such as herbivore living and operating on the same surface of the same habitat .

Once a new physical design was achieved most of the evolutionary changes that took place where focused on improving the EROEI and survivability by either one or by the combination of cognitive evolution for developing cognitive strategies and physical evolution to achieve advantageous position with size, strength, agility or others.

The genetic assets of a species can suggest the spectrum of evolutionary routs

Although the energetic balance state and trend of a species and its evolutionary trend are correlated to the energetic states and trends in its ecological community, the species spectrum or the genetic potential of the possible evolutionary rout of morphological process, is determined by the genetics factors or the genotype of an organism:

Genotype: is the part of the DNA sequence responsible for the genetic makeup of a cell, and therefore of an organism or individual, which determines a specific characteristic (phenotype) of that cell/organism/individual.

Genotype is one of three factors that determine phenotype, the second factor is the inherited arsenal of traits from earlier evolutionary adaptation of bodily and cognitive modules that are rudimentary/dormant or that are currently in use for other tasks but can be updated, modified or converted to perform the new needed tasks to overcome new conditions, and the third is the non-inherited environmental (abiotic and community of life) factors.

Phenotype: is the composite of an organism’s observable characteristics or traits, such as its morphology (the change in the form and structure of organisms and in their specific structural features), development, biochemical or physiological properties, behavior, and products of behavior. A phenotype results from the expression of an organism’s genetic code, its genotype, as well as the influence of environmental factors and the interactions between the two. When two or more clearly different phenotypes exist in the same population of a species, the species is called polymorphic.

Tracking species’ states and evolutionary trends

To track an evolutionary process of a long gone extinct species based on ancient abiotic factors data, genetic data, limited fossil record and living relatives that have evolved further from the ancestors, demands research methods that are more holistic and which are based on methods used in theoretical physics that are utilizing the principals of thermodynamics to assess and model biological systems by assessing the energetic balance and the state of individuals organisms and species such as the energy stored as mass, the stock of potential work energy (ability to function) and the EROEI of activities all in the context of the energy stocks, flow and exchange between the entire web of life and its interaction with the energetic apparatus of the environmental systems of abiotic factors.

The survivability potential of an organism and a species can be delineated by its energetic efficiency and it is measured by the Energy Returned on Energy Invested (EROEI) and thus can delineated the organisms’ evolutionary state and trends.

Biological systems are systems of energy acquisition and storage; the exchange of energy within and between biological systems and their environment is the engine of life and evolution. The exchange of energy within and between biological systems can be defined by the specification of such activities, living organisms perform many activities; some are internal and rudimentary, and some are external or behavioral: all the activities performed by Individual biological systems can be grouped into two fundamental categories:

  • Energy input oriented activities
  • Energy output oriented activities

The balance of these two activities is measured in EROEI (Energy Return (input) On Energy Invested (input)) and in the accumulated energy stock measured by metabolized calories stored in the mass of organisms.

The measure of the states and trends of specie’s individuals, classes and groups can predict the evolutionary direction of a species:

We can observe the emerge of evolutionary trends in a species by measuring the short term and lifetime EROEI of their input activities and the short term levels of the stocks of working energy that is stored in the mass of individuals in accordance to their specific energetic need during each stage in their life histories and developmental stages (e.g. growing stages or adulthood), we can then compare to previous generations of individuals to identify generational changes in species energetic balance and stocks that suggest an evolutionary trends.

Measuring concurrent states and short term trends of specie’s individuals/classes/groups

Daily EROEI (short term state): measures the concurrent state of individual organism and groups of individual organisms of same species: measured by deducting the amount of energy output from the amount of energy input during the term of one planetary rotational cycle of 24 hours which includes all the daily activities as in the case of primates, or by averaging another short term cycle such as metabolic cycle, seasonal cycle etc. as in the case of other organism:

  • If the resulting daily EROEI is negative than the individual is in a state of short term energetic deficit
  • If the resulting daily EROEI is positive than the individual is in a state of short term energetic surplus
  • If the resulting daily EROEI is equalized than the individual is in a state of short term energetic equilibrium

Daily energy stock balance (short term trend): measures the short term trend in the energetic stock of individual organism and groups of individual organisms of same species: calculated by adding the daily EROEI at the end of the daily cycle to the stocks of energy (that was stored in the mass of individual organism, at the beginning of the daily cycle (or other short term cycle):

  • If the resulting daily energy stock balance is negative than the individual’s short term trend is of recession
  • If the resulting daily energy stock balance is positive than the individual’s short term trend is of affluent
  • If the resulting daily energy stock balance is equalized than the individual’s short term trend is of equilibrium

Any change in the environmental conditions in the organism’s habitat will change the energetic balance between the daily activities and will affect its long term survivability and the generational evolutionary trends of its population (and species). Monitoring the short term level of stocks of energy of individuals within a species, can highlight the points of transition between positive, equalized and negative energetic balance which in term can indicate short term trends as well as change in the direction of long term evolutionary trends in a species and can indicate changes in species fitness strategies.

Measuring longer term states and trends of specie’s individuals/classes/groups at a defined developmental stage or during defined periods of time

Longer term EROEI trend – measuring the success of the behavioral part of the fitness strategy during the span of defined period of time or life histories stage (adolescents, adulthood etc.) of an individual organism and groups of individual organisms of same species: calculated by factoring in a matrix all the daily EROEI cycles values during the span of longer term periods of weeks, months and years:

  • If the resulting longer term EROEI trend is negative than the individual/group/species was/is in a chronic energetic deficit during the span of the defined period
  • If the resulting longer term EROEI trend is positive than the individual/group/species was/is in a constant or a developing energetic surplus during the span of the defined period
  • If the resulting longer term EROEI trend is equalized than the individual/group/species was/is in an energetic equilibrium during the span of the defined period

Longer term energy stock balance – measuring the success of the physiological part of the fitness strategy during the span of defined period of time or life histories stage of an in individual organism and groups of individual organisms of same species: calculated by factoring in a matrix all the daily stock balance values during the span of longer term periods of weeks, months and years:

  • If the resulting trend of all daily energy stock balance values is negative than the trend for individual/group/species body mass in the defined period of time is of contraction
  • If the resulting trend of all daily energy stock balance values is positive than the trend for individual/group/species body mass in the defined period of time is of development
  • If the resulting trend of all daily energy stock balance values is equalized than the trend for individual/group/species body mass in the defined period of time is of stagnation

By measuring longer terms cycles/period of monthly, yearly and seasonal energetic balance in individuals, groups and population by deducting the stock of stored energy at end of a cycle/period from the stocked energy at the beginning of a cycle to detect transitional foresees in the energetic balance between the species and its environment.

Measuring lifetime state and trends of a generation of specie’s individuals/classes/groups by lifetime EROEI trend and energy stock balance

Lifetime EROEI trend (from sexual maturity to death) – measure the aftermath success of the behavioral part of the fitness strategy of the past generations of an individual organism and groups of individual organisms of same species: calculated by factoring in a matrix all the daily EROEI cycles values during the span of an organism lifetime:

  • If the resulting lifetime EROEI trend is negative than the individual/group/species was/were in a chronic energetic deficit and it suggest a failing behavioral fitness strategy
  • If the resulting lifetime EROEI trend is positive than the individual/group/species was/were in a constant or developing energetic surplus and it suggest a development of a successful behavioral fitness strategy
  • If the resulting lifetime EROEI trend is equalized than the individual/group/species was/were in an energetic equilibrium and it suggest a continuum of an optimal behavioral fitness strategy

Lifetime energy stock balance (from sexual maturity to death) – prospect the morphological trends of the physiological part of the fitness strategy in the next generation of in individual organism and groups of individual organisms of same species: calculated by factoring in a matrix all the daily energetic balance values during the span of an organism lifetime:

  • If the resulting lifetime trend of all daily energy stock balance values in individual/group/species was/were in a decline than the prospect of morphological contraction in the next generation of individual/group/species is more probable
  • If the resulting lifetime trend of all daily energy stock balance values in individual/group/species was/were is in developing surplus than the prospects of morphological development in the next generation of individual/group/species is more probable
  • If the resulting lifetime trend of all daily energy stock balance values in individual/group/species was/were equalized than the trend for individual/group/species is physiological stagnation in the next generation is more probable

The stored energy in each level of the web of life and the flow and exchange of energy between biological systems and environmental systems can describe the properties of the community of life and can identify their interactions with each other at every levels of the food chain and can be describe the behaviors or fitness strategies of an organism.

Evolutionary processes fundamentals

The most important tasks of individuals of a species are eating, mating and reproduce in a way that is ensuring the survival of the minimum number of offspring’s that are required for growing the population size in times of development or recession in the ecosystem; when an ecosystem reaches the state of climax community the reproduction requirement become less demanding; just enough offspring for maintaining the population size.

When the energetic balance is drawn to the negative side due to recession in the ecosystem (which ignite competition for diminishing resources and habitat) or introduction of a threat initially the main strategy focus shift into minimalizing the energy and time invested in regular less urgent activities such socializing to compensate for the time and energy invested in the important activities or waste of time and energy passively consumed by the enforcement of unproductive behavioral activities such as fighting, hiding, escaping etc.

Every activity other than the necessity can jeopardize animal’s energetic balance constant which result in a negative EROEI and are not directly contributing for the process of energy consumption or reproduction or comes on the account of to the physical fitness supporting activities of consumption (foraging for food and eating, sleeping) and activities supporting wellbeing (sleeping, playing, socializing etc.).

These activities are all related for a non-equilibrium state of a species and a non-equilibrium state of its environment (a state of disturbance or a state of succession) this activities are all related for Intraspecies and interspecies competition (resources, space, mating) and basic survival rudiments (not being injured or killed).

Every strategy used or developed in order to address this kind of negative energetic state is considered a non-equilibrium strategy:
Social strategies are usually the result of a non-equilibrium state of a species.

Any type of social organization that consist outside the period of mating and parenting or that include more individuals in the group in excess of a core family (one male and one female and may include dependent juvenile and adolescent offspring)

Individualistic strategies are usually the result of a state of equilibrium of a species

Without the need to develop social strategies an animal will invest in the individuals of the species to ensure both genders will have maximum level of equilibrium, it means that both genders have equal or similar size, weight and physiology as a constant static state throughout the individuals life histories that will include maximum longevity and maximal lifespan which can last for many generations for as long as the environmental condition maintained.

The highest level of energetic equilibrium that can be reached by a species is the ability to maintain consistent average of positive daily EROEI throughout its total lifespan. Only a species that can maintain the highest level of energetic equilibrium for long periods can reach the maximum extended longevity, lifespan and fitness throughout its life histories.

Positive non-equilibrium evolutionary process of adaptation to improve the EROEI by consuming more energy:

Expansion of primary producer’s biomass, biodiversity and energetic values in the form of regular supply and availability of energetic products driving the habitat ecosystem towards equilibrium by forming parallel diversifying evolutionary processes in the communities of organism on the path to climax community. New traits are developing, maintained and improved.

Negative non-equilibrium evolutionary process of adaptation to improve the EROEI by reducing energy expenditures:

Recession of the primary producers’ results in diminishing biomass, biodiversity and energetic products (leafs, fruits etc.) and services (shelters, habitat) are negatively disturbing the EROEI and energetic balance of the consumers.

Consumers in respond will return to old traits as their more recent strategies which are based on latest traits start to fail in supplying the sufficient EROEI that previously supported such more energy expensive traits as well as the development of new traits.
Consumers will return to strategies of energy conservation which are antagonistic to developmental stages or counter developmental evolution i.e. regression. Energy expensive organs parts may also be in the brain where cognitive traits reside, and a “cognitive recession” will occur.

The quality of cognitive traits is depending upon the ancestral heritage of brain part and concurrent assets of functionality circuits.
Brain parts and modules can diminish, synaptic rearranged or completely repurpose when past biomechanical strategies which supported more agile locomotion strategies which demanded higher “cognitive infrastructure” for higher real-time processing become old cognitive infrastructure for older strategies based on previous ancestral adaptations which rendered obsolete as conditions changed (e.g. the brain of a bipedal primate traveling and navigating in speeds of 10-20K/h on the two dimensional surface does not need the cognitive agility of a ricochetal brachiation traveling and navigating in speeds of more than 50K/h in the three dimensional environment of the forest canopy).

 

Fitness strategy – the behavioral output of a physical system with a measurable or observable manifestations in organism’s internal and external systems of an evolutionary trend of adaptation

Fitness is the driver of all strategies and it can describe as the long term state of health and well-being of an organism to be better positioned for reproduction. Fitness is the ability of an organism to perform all aspects of daily activities by supporting the energetic expenditure while maintaining constant long run average of positive or equalized energetic balance in accordance to the animal’s life histories of developmental stages and ultimately successfully achieving the fulfillment of the most important task of living organism – reproduction. Fitness can only be achieved if the short term and long term EROEI and energetic balance of an organism are equalized in the end of the long term cycles and life histories stages.

Fitness strategy is the sum of all phenotypic strategies of physiological systems and behavioral tactics of inherit and learned traits and skills, that evolved as a result of incremental genetic mutations over generations of individual organisms that posteriori were beneficial for the processes of sexual selection, reproduction and for improving the survivability of their offspring’s which ultimately improved some individual organisms fitness which advance their reproduction and their genetically privileged offspring reproduction resulting in evolving homogenous genetics lineage adapting and multiplying and once they reach an equilibrium of individual fitness and stable population in the boundaries of their habitat carrying capacity they become a species.

Behavior: defines the range of actions and mannerisms made by individual and systems of organisms in conjunction with themselves or their environment, which includes their internal systems interactions (the cognitive system, the biochemistry in it cells, the biomechanics etc.) and their interaction with other intraspecies and interspecies systems of organisms and with the physical and structural environment and the abiotic factors. It is the response of an organism to various stimuli or inputs, whether internal or external, conscious or subconscious, overt or covert, and voluntary or involuntary.

Strategy: is an underlying rule for making decisions about a certain behavior, strategy provides an animal with a set of tactics that are adaptive in various circumstances. A tactic is the action that is taken by an animal in order to achieve a specific goal. For example, an animal encounters an obstacle and its strategy is defined by two tactics that may allow the animal to pass the obstacle: A. jump over it, B. crawl under it.

Considering the environmental conditions, the surroundings, and the size of the obstacle, the animal will decide between the two tactics dictated by its strategy. In the context of a mating system, this means that individuals in a given population have strategies that allow them to obtain mates in different ways to maximize their reproductive success given their phenotypic, environmental, or social circumstances. There are different types of strategies such as:

  • Dynamic strategies: are the aftermath of the adaptive responses of a species for constantly changing or unstable environmental, interspecies and intraspecies factors and conditions (such as in the case of responding to a recessional or developmental process in their ecosystem). Dynamic strategies may be considered Mendelian, developmental, conditional, or a combination of them:
  • Mendelian strategy: depends on a genetically determined phenotypic difference, such as body size and physiological modules and parts
  • Developmentally driven strategy: is associated with phenotypic differences caused by varying conditions during the course of development that affect physiological modules and parts.
  • Conditional behavior strategy: depends not on the genetic or developmental impact, but on external factors. These may include the cognitive ability to combine or sequence an arsenal of tactics under one behavior or a set of behaviors, the number and the sophistication of the sets and combinations of tactics which are employed in response for various situations and their success will be determine by the range of cognitive modularity, capacity and flexibility of an animal to calculatedly choose a proper more fine-tuned behavior in order to interact and react to a wider range of situations and circumstances with the lowest energetic expenditure and risks as possible, and this range of tactics form the spectrum of behavioral responses and the cognitive ability to invent, combine and memorize several tactics to produce new behaviors define the intelligence of an animal.

Stable strategy: the end result of a successful fitness strategy and it will occur when constant fitness of species has been achieved and it persists for as long as the environmental conditions prevail. Stable strategy represents stagnation in the process of evolutionary adaptation of a species and it is correlates to the state of a climax community or an ecosystem in a state of equilibrium.

In a state of equilibrium every organism and species conditional behavior strategies are adjusted and fine-tuned for optimizing its own energetic balance which minimize its impact on the community’s energy stocks and support the equalization of the total exchange and flow of energy in an ecosystem.

In a climax community the energetic needs of all the community species are at the optimal energetic balance or the minimal state of EROEI that is needed in order to maintain the fitness of organisms and species.

Species in a climax community maintain their population numbers and limit their energetic consumption at optimal energetic balance in accordance to their life histories stage and the optimal EROEI of the short term and long term cycles (daily activities, reproduction cycles etc.).

Fitness strategy defines the main trend in a species evolutionary process (it’s include organism’s biological and cognitive systems) and the direction of the development which aimed to achieve a state of fitness equilibrium by overcoming (adapting) the environmental challenges which affect their short term energetic balance and EROEI in a negative way.

Individualistic strategies in mammals are usually considered as R-selection species and group strategies are usually considered as r-selection species

Fitness can be improved in several ways depending on the urgency of the condition and the effect it have on reproduction and on the availability of genetic potential, it can be an adaptation for gradual or sudden changes in the environmental conditions or relative to certain condition regardless to the state of the ecosystem (such interspecies competition).

Phenotypic changes are incremental changes over generation in the responsible genes which survived after generations of being the favored traits for sexual selection (or by pure chance); there are few ways phenotypic changes occur:

  • By improvement of currently active traits – for example: a decrease in the mass and strength of certain groups of muscles and bones favored by sexual selection or in response to other conditions.
  • By moving forward a recently retrogressed trait – for example: an increase in the mass and strength of certain groups of muscles and bones favored by sexual selection in response to increasing intraspecies or interspecies aggression.
  • By “awakening” a retro-trait that was “dormant” for a very long time to perform the same tasks it was design for that once again has become useful for the animal fitness in the face of a new condition.
  • By transforming a trait to perform completely different tasks then it was design for in the face of a sudden new condition.

For example the process in which a species that have lost its tail due to energy conservation strategy is easily achieved but the opposite process of growing back a tail that is fully functional will take longer to achieve.

The process of a change in the properties and behaviors of organism’s systems and their subsystems over generations of sexual reproduction indicates, post factum, the strategies that drove a species evolutionary rout of behavioral (cognitive) and/or morphological (physical) adaptations which ultimately improved such species fitness.

Fitness strategy is the sum of organisms’ behavioral strategies and the physical properties of its internal systems

Fitness strategy can be defined and grouped by the functions of their behavioral cognitive output in the form of strategies which sometime incorporating internal subsystems of modules and parts that participate in and control all the physiological and cognitive activities:

Feeding strategy: determine and define the behavioral output of the energy acquisition process of set of activities and tactics that in motile animals that include foraging for items of stocked energy (food) and ends when the items of stocked energy are within the reach of the digestive systems (mouth) which is the entry point for the metabolic system.

Feeding strategy defined by and describes the systems of energetic consumption by individual organism within a species it include the following systems:

Energy metabolism, stocking and energy distribution systems: autonomic and rudimentary systems for utilizing the consumed objects of energy by processing (digesting), regulating energy utilization and outflow, stocking energy and distribution of the energy to the different systems that are involved and various activities and tasks supporting the organs, parts and modules that are included in the participation of all other activities and tasks which require metabolized work energy available for output for all the behaviors which are included in the fitness strategy.

Metabolism: the sum of all chemical reactions that occur in living organisms, including digestion and the transport of substances into and between different cells, in which case the set of reactions within the cells is called intermediary metabolism or intermediate metabolism.

Metabolism have three main purposes: the conversion of stocks of consumed energy (food) to work energy in order to run cellular processes, the conversion of food to building blocks for proteins, lipids, nucleic acids, and some carbohydrates, and the elimination of nitrogenous wastes.

These enzyme-catalyzed reactions allow organisms to grow and reproduce, maintain their structures, and to respond to their environments. Metabolism is usually divided into two categories:

  • Catabolism: the breaking down of organic matter, for example, by cellular respiration.
  • Anabolism: the building up of components of cells such as proteins and nucleic acids.

The chemical reactions of metabolism are organized into metabolic pathways, in which one chemical is transformed through a series of steps into another chemical, by a sequence of enzymes. Enzymes are crucial to metabolism because they allow organisms to drive desirable reactions that require energy that will not occur by itself, by coupling them to spontaneous reactions that release energy. Enzymes act as catalysts that allow the reactions to proceed more rapidly. Enzymes also allow the regulation of metabolic pathways in response to changes in the cell’s environment or to signals from other cells.

The metabolic system of a particular organism determines which substances it will find nutritious and which poisonous. The speed of metabolism, the metabolic rate, influences how much energy (food) an organism will require, and also affects how it is able to obtain that food.

One of the features of metabolism is the similarity of the basic metabolic pathways and components between even vastly different species. For example, the set of carboxylic acids (the intermediates in the citric acid cycle) are present in all known organisms, being found in species as diverse as the unicellular bacterium Escherichia coli and huge multicellular organisms like elephants. These striking similarities in metabolic pathways are likely due to their early appearance in evolutionary history, and their retention because of their efficacy. The metabolic system includes the following systems and their parts:

  • Digestive system: digestion and processing food with salivary glands, esophagus, stomach, liver, gallbladder, pancreas, intestines, rectum and anus.
  • Urinary system: kidneys, ureters, bladder and urethra involved in fluid balance, electrolyte balance and excretion of urine.

Locomotion strategy: is the behavioral output of a motile organism and it includes all the systems involved in the motor control and the mechanisms which responsible for the execution and the performance of motile activities such as walking, running, climbing, swimming, brachiation etc. intended for performing all type of movement required by the feeding and reproduction strategies. The locomotion systems and parts include skeleton, muscles, nerves, blood systems etc.:

  • Muscular system: allows for manipulation of the environment, provides locomotion, maintains posture, and produces heat. Includes skeletal muscles, smooth muscles and cardiac muscle.
  • Skeletal system: structural support and protection with bones, cartilage, ligaments and tendons

Reproduction strategy: behavioral and physiological systems that are involved in and supporting the activity of duplicating the genetics of males and females from one generation of organism to the next generations and for ensuring the survival of the offspring’s, the activities of reproduction includes courtship, engagement, sexual interaction and intercourse, and the different sets of activities performed by the breeding animals for ensuring the survival of their offspring’s such as mass reproduction, single parent, coupling, nourishing, protection, education etc. The reproductive system includes the following systems and their parts:

  • Reproductive system: the sex organs, such as ovaries, fallopian tubes, uterus, vagina, mammary glands, testes, vas deferens, seminal vesicles and prostate

Operating systems (neuroendocrine system): the management of all the organisms’ subsystem and it is in charge of the coordination and operation of all subsystems which incorporated all the behavioral output of the fitness strategy, the operating system includes autonomic and cognitive systems:

  • Endocrine system: communication within the body using hormones made by endocrine glands such as the hypothalamus, pituitary gland, pineal body or pineal gland, thyroid, parathyroid and adrenals, i.e. adrenal glands
  • Nervous system: collecting, transferring and processing information with brain, spinal cord and peripheral nervous system.

Other systems of distribution of energy, oxygen and maintenance and repair systems:

  • Cardiovascular or Circulatory system: pumping and channeling blood to and from the body and lungs with heart, blood and blood vessels.
  • Respiratory system: the organs used for breathing, the pharynx, larynx, bronchi, lungs and diaphragm
  • Immune system: protects the organism from foreign bodies
  • Lymphatic system: structures involved in the transfer of lymph between tissues and the blood stream; includes the lymph and the nodes and vessels. The lymphatic system includes functions including immune responses and development of antibodies.
  • Integumentary system: skin, hair, fat, and nails.
The cognitive system the facilitator and operator of the locomotion which is the determine factor of the fitness strategies

The cognitive system (“cognition”) has two main subsystems: system of controlled cognition for motile decisions and system of rudimentary cognition for automation of biomechanical sequences as part of locomotion and movement and for immediate and instinctual responses:

The cognition is the operating system of biological systems: it is based on neurological tissue of modules as hardware and it is measured by the input, processing and output capacity and by the velocity of data flow and threading which are determined by the arrangement, amount, type and bandwidth of neurons, synaptic connections, wiring and brain modules as well as the pathways in which information flow.

The cognition evolved to answer the needs of the feeding strategy and the preservation of metastability by operating the biomechanics modules in accordance to motile decisions that and to execute the locomotion in accordance to the fitness strategy.

Cognitive abilities are brain-based skills intended to carry out any task from the simplest to the most complex:

They have more to do with the mechanisms of how we operate via logic and how we learn: remember tactics, solving problems, and pay attention that can be compared to wisdom, rather than relay on a database of tactics and symbolic triggers that are based on memorizing experiences and theoretical non-personal experience which can be compared to knowledge.

The performance of the cognition will be determined by the environmental challenges and opportunities that drove their development: the structural complexity of the environment, the distribution, location and properties of objects of energy that are suitable for their current digestive system, the sophistication of the species biomechanical hardware such as the motoric system, these will define the type of cognitive calculations that are needed to support the execution of the locomotion, the type of EROEI strategy of passivity (relayed less on cognitive power) or agility (demands more cognitive power especially when the agility pushed to the extremes of speed and velocity), inherit brain modules and cognitive systems, intraspecies and interspecies interactions (social and threats) and many more other determining factors.

The cognitive initiation, control and operation of locomotion by an intentional agent with motile objectives, is only possible by using energetic expenditure from an internally stored stock of energy. The energetic stock is metabolized and distributed to the motoric system where it is utilized for performing biomechanics sequences which results in mobilization and traveling for achieving motile objectives (move away from a current location (point A) to another location (point B).

Locomotion is the result of motile decision to answer metastability objective (feed, drink, reproduce, survive threats and hazards etc.) and it is driven by the need to mobilize to another location where it safe, important or where consumable object of energy is located.

Agile locomotion strategy: means faster locomotion that includes faster gait speeds and rapid changes of route and utilization of trajectories and is the result of a sufficient surplus energy that is the result of positive EROEI from the feeding strategy which is sufficient enough to support all the daily activities which maintain the life of an organism including the activities involved in passing its genes to the next generation which in whole defines the species fitness strategy. Over time in incremental of generations the agile locomotion strategy can improved and maintained by focusing on prime energetic objects for growing the surplus balance by increasing the energetic input of higher density, intensity of nutrients, improving metabolism or increasing the amount of the consumed energy stocks by improving the biomechanics functions and EROEI efficiency or by improving the current motile decision making mechanism for improving the yields of the feeding strategy by enhancing the capacity and functionality of existing cognitive and physical systems and modules and by developing new ones.

Passive locomotion strategy: means slower locomotion that includes slower gait speeds and less changes of route or use of trajectories and is the result of an energy deficit resulting from environmental conditions that impose negative EROEI on the current feeding strategy due to increase in foraging time due added distance, from other added activities such as avoiding hazards and treats, as a result of increase in competition and reduction of yield due to abiotic conditions and others which render the current feeding strategy as inefficient and insufficient for supporting the previously considered regular energetic expenditure of the regular daily activities. Over time in incremental of generations the passive locomotion strategy will balance the deficit by adjusting the animal to conservation based fitness strategy based on the availability of energy by reduction of the animal own mass of energetic stock, eliminate energy expensive physical parts and modules (like the tail that is not needed when moving slow)

A passive locomotion strategy can also work to eliminate energy expensive behaviors for example socializing activities by reducing the energetic toll of fights by avoiding other species and even by changing the feeding and reproduction strategies by transforming the interspecies competition for mating, territory and food from an energy expensive and risky physical interaction into symbolic ritual of passive aggressive aggression and intimidation via virtue signaling utilizing gestures, vocals and other communication memes or by crating standards of hierarchal access classes determine by certain properties, features and means that classify each of the contenders in a certain class or status according  to a general or culturally excepted “stud magnitude” scale which measure their relative position to each other by assessing and comparing their possessions in terms of physical and behavioral assets (size, strength, shape, color, specific body part, behavioral performance etc.) or in comparison to the “standard” of the general population of the species, related to their gender and life history stage.

In order for a motile organism to survive and pass it genes, it must be able to accomplish and complete the daily cycle of energy input that performed by the locomotion and feeding strategies (which includes traveling, locating, acquiring, consuming and metabolizing objects of stored energy) with a positive and sufficient EROEI; which means that the energy output invested in locomotion and digestion must return constant surplus of energy from the daily consumption of such objects of stored energy, that is sufficient for the energetic expenditures of all other daily activities.

The locomotion strategy brings specific objects of energy that are suitable for process by the organism’s specific type of digestive system into the reach of its opening  and in accordance to the feeding strategy which determine the choice of energy object types that will be consumed.

Locomotion strategy must support this fundamental when performing a locomotion trip from point A to point B where consumable object of energy is located.

The type of locomotion of an animal may determine or be determined by its interest in a certain type of objects of energy and the distribution and location of the stocks and their certain EROEI value that is compose of multiple factors such as the object energetic intensity, density and quantity and the energetic expenditure which are involved in the processes of it acquisition compared to its metabolism EROEI  its nutritional values and the concentration and ratio of nutritional necessities in terms of the amount, variety and intensity of carbohydrate which provides metabolic work energy, lipids (fats and oils) provide stockable high intensity energy for future metabolism or for thermoregulation, proteins for growth and repair of cellular tissues, minerals and vitamins to perform hundreds of roles in biological systems such as maintain the shore up bones, heal wounds, and bolster the immune system, to convert food into energy, and repair cellular damage.

Reproduction strategies fundamentals and their dynamic feedbacks with the locomotion and feeding strategies

There are two main types of reproduction strategies (that can be described as antagonistic – meaning that when a species is reliant on one it use less of the other) that are in a direct relation with the genetic assets of an organism, means they determine by the organism’s physical brain properties and cognitive capacity which determine the spectrum and levels of behavioral strategies and instrument it can accommodate.

Two animals may have similar physical strategies (shark and orca) which are determine by the efficiency of their locomotion but the difference between their cognitive systems will determine their reproduction strategy.

These two types of strategies are referred to as K strategy and the r strategy; K strategists are referred to as “K-selected” and r strategists are referred to as “r-selected.”

All living organisms are supposed to fall somewhere on this continuum between the two extremes (r and K). Also, organisms can be differentiated from one another in terms of their relative reliance on one strategy over the other.

r ____._________._________.______.________.__________._________._________._______K

bacteria mollusks insects fish amphibians reptiles mammals primates

  • r-selection species requiring small energy investment to produce many “cheap” offspring and live in unstable environments.
  • K-selection species requiring high energy investment that produce few “expensive” offspring and live in stable environments.
K-selection apex species

A group of animals can be referred to as a species only if it reaches energetic equilibrium at any trophic position in the community of life and it will then be considered to be an outgroup species

In a non-equilibrium environment species will be in a developmental state or a state of transition and it will then be considered to be a sister group of the outgroup species until it reaches equilibrium:

Apex species

Apex carnivore consumer species can be r-selected or K-selected: is a carnivore species residing at the top of a food chain upon which no other creatures prey. Apex predators are usually defined in terms of trophic dynamics, meaning that apex-predator species occupy the highest trophic level or levels and play a crucial role in maintaining the health of their ecosystems.

Apex herbivore consumer species can only be K-selected: is defined as one only if it reaches equilibrium at the highest trophic position possible: An Apex Consumer is a non-predatory species that have no intraspecies competitors/predators and therefore position at the top of the food chain of its climax community (the carrying capacity of a biological species in an environment is at optimal population size which the environment, if the supporting condition persists, can sustain indefinitely, given the food, habitat, water, and other necessities available in the environment).

A species that reach the level of Apex Consumer/will include many of the following characteristics and features of a K-selection species (e.g. r/K selection theory):

  • The species size, weight, morphology and physiology will reach a constant static state
  • The species lifespan and longevity are extended to the maximum
  • The species as whole and most individuals will maintain consistent level of positive EROEI short-term (weekly/monthly) and long-term (yearly/lifetime).
  • Reach and maintain an optimal size for maintaining the surplus (largest size possible for an organism while maintaining the maximal state of equilibrium)
  • Energy used to make each individual is high
  • Few offspring are produced
  • Late maturity, often after a prolonged period of parental care
  • Long life expectancy
  • Individuals can reproduce more than once in their lifetime
  • Most individuals live near to the maximum lifespan

K-selected species have a relatively long life span, produce relatively few offspring, the offspring have low mortality rates and the parents provide extensive parental care.
The offspring are also relatively intelligent so that they can internalize the lessons from their parents.
Primates are the most K-selected because their young are truly helpless- they necessitate years and decades of parental care and tutelage and the parents usually only produce one offspring at a time.

The range of life histories stages, periods and developmental stages are corresponding with the survivability and longevity of the individuals of the species:

  • K strategy: long physical and cognitive developmental period – survivorship is high, most mortality later in life span
  • In between: constant mortality throughout lifespan
  • r strategy: : short physical and cognitive developmental period, high mortality early in life, low survivorship into late life span

K-selected species have to limit their body size to the optimal energetic efficiency that can be achieved by a physiological design in relation to the medium of the locomotion (liquid, branches, ground etc.) and the biomechanical tools and technics of locomotion (fins and jet propulsion in liquid, quadruple or bipedal of arboreal locomotion of vertical gripping and climbing, walking/leaping on top or swinging/ricocheting from the bottom of branches etc.).

Another considerable factor is the carrying capacity of their niche in the ecosystem in term of the energetic state and trends and their effect on consumable energetic products available for them:

For example growing in body size when the locomotion strategy is brachiation and the medium of movement is trees’ branches will eventually force the species to move to the ground due to the thermodynamic limitations of their locomotion strategy in term of EROEI and the physical limitation of the structural infrastructure of branches to support their weight (i.e. gorilla), which will effect negatively the EROEI of the species feeding strategy due to the higher energetic expenditures of quadruple knuckle-walking locomotion compare to brachiation which they will have to compensate by different strategies limited by the type, range and availability of energetic product that they can consume and metabolize:

When larger animal is not changing the diet then it will have to compensates the higher energetic needs by increasing the energetic efficiency of the current consumable products (i.e. fruits, leafs etc.) they can achieve that also by behavioral adaptation such as increasing the amount of consumable products, increasing the foraging range and investing more time in consuming, focusing on products with higher energetic intensity in the same category and in the same metabolic range (i.e. eating young leaves instead of mature leaves) and widen the range of the consumable products limited only by their physical abilities and properties (i.e. jaws, teeth etc.)

They can achieve larger size in the longer run by morphological adaption process, an adaption for increasing the metabolic energetic input capacity and EROEI of the feeding strategy by developing more energetically efficient metabolism or higher metabolic capacity per feeding session, they can also develop new strategies for walking on the ground which will improve the EROEI of the quadruple knuckle-walking locomotion by adapting a strategy that is more efficient when walking on top of surfaces instead of gripping structures such as bipedal locomotion all depends on the food sources in their habitat and their daily necessities.

They can also adopt a strategy of energetic expenditure redistribution by decreasing the energetic expenditure of other activities (socializing, sleeping etc.) by reducing the time invested in them or by eliminating them, they can also try to reduce the time involved in the process of consumption (foraging) of the current consumable products if they can develop a behavioral strategies to energetically support a process of increasing in body size for example stockpiling energetic resources when they abundant (if conditions or type of food resource deem it possible i.e. nuts and grains, tree bark and even meet in winter freeze time) and warehousing them in underground pantry or they can go through a longer term morphologic adaption for stocking metabolic energy in fat, they can develop behavioral adaption of stealing resources from others and more.

Another way to support bigger physiology is changing or expanding the metabolic focus or range to include energy sources with higher energetic intensity such as meat but that presents new challenges when the feeding strategy is changed from vegetarian (i.e. frugivore, herbivore etc.) to omnivore or carnivore diet, metabolizing meat do not need such a radical adaptation in mammals as the need for digesting high concentrated protein in the form of mammary gland milk during their initial post-birth developmental stage makes their digestive system more adequate for digesting meet and many primates are more of an omnivore as many of them include insects as a food source, and although the metabolic adaption to digest meat is possible without any physiological adaptation, getting the meat and eating the meat demands certain traits of hunting and consumption which require morphological adaption of developing the weaponry of predation or the development of new behavioral strategies to overcome physiological limitation such as hunting in groups.

For r-selected species, the body is growing much faster so the “small copies” catch up with the physical development (faster growth rate) r-selection species do not have the maximal size limitations which are based on the size of the brain (in term of volume and weight) and the energetic expenditures which are involved in it developmental stage and depending on the medium (animals living in liquid do not have the brain size limitation due to the lack of gravitational factors (i.e. whales)) and they can grow larger in mass and weight bound only by the energetically limitations of their environment and one of the advantages of larger size is that they can have more offspring which they can produce with lower amount energy invested in each.

The different strategies to achieve higher energetic input in order to support larger size are based on physical development and are usually related to the ecosystem state as the morphological adaptation of growing in mass demand a continuum of energetic surplus which is normally related to an ecosystem in stat of development or equilibrium.

Other strategies are contrary to growth and are related to energy conservation and are correlated to species in energetic deficit due to competition or other community of life factors are normally related to an ecosystem in recession due to abiotic or other environmental conditions.

The developmental limitation of a large brain in K-selection species

The larger brain that is needed in order to facilitate higher cognitive behaviors sets a cap for the size of the offspring at birth: due to a physiological barrier govern by the practical limitations which arises from the pelvis roll in locomotion, limiting the pre-birth tissue and skeletal expansion capacity to a specific size of brain which unlike other body organs needs longer time to develop the neurologic complexity and the mass involved in advanced fitness strategies and it have to come with proportional body mass to support its biological needs, which remove the option of having miniature copies.

Brain size limitations on EROEI and developmental physiology pre/post-birth

The limitation on the size of the brain at birth is governing the fitness strategy of more cognitive developed animals for maximizing the pre-birth development capacity to shorten (as physiologically possible) the post-birth period of dependency and custody. The long post-birth and pre-birth developmental process presenting a large energetic toll on the parenting adults: the energetic investment in the developmental period of the offspring is determine by the ability of the species to maintain high energetic surplus for the entire process and therefore needs energetically effective feeding and locomotion strategies to support higher levels of EROEI and long term energetic surplus.

Thermodynamic and functional mechanic limitations of large brain

Another limitation of the brain weight for surface quadruped (as most mammals are) is in the pronograde (horizontal body position) design and arrangement and distribution of the body parts between the front and the rear for optimizing the EROEI and the agility by weight equalizing and motile supporting balance: in quadrupeds the optimal design is a rectangular shape of the motor system where the limbs forming a vertical diagonals and the top of the back and the ground forming the horizontal diagonals, together with the flexible spring like spine and the triple springs systems of the limbs it is the optimal design for high performance or for optimal EROEI which is related to the gait strategy which drive the optimization of the locomotion to a certain purpose of energetic preservation due to low returns of the motile tasks (e.g. to cover long distance with the most efficient locomotion EROEI for energetic preservation as camels) or to invest high bursts of energetic output at times in order to obtain much higher energetic returns from the costly energetic expenditure of the gait, for example to accelerate fast and to be able to change direction in order to catch a high energy protein meal or to escape becoming that meal (e.g. Chita & gazelles), to maintain high speeds (i.e. stamina) for long distance (e.g. wolf & deer) etc.

The pronograde quadrupeds was designed for the minimal stable support (as a table) of an horizontal mass to center of gravity it is the optimal method for traveling on horizontal surfaces and it is easily adapted for optimizing energetic efficiency or for maximizing agility and performance and the distribution of weight between the rear and the front and the length and flexibility of the spine and limbs are key factor for the locomotion performance and EROEI of quadrupeds.

The energetic expenditure of quadruped designs are a factor of the locomotion strategy drivers and generally quadrupeds that developed proportionally longer limbs with short torsos and less flexible spine springs are better for reducing the locomotion EROEI (they compensate the limitation of a short spine spring and the lack of the balancing tail by extending the their limbs springs and reducing the size and weight of their heads) while quadrupeds shorter limbs and longer back spring are much more energetically expensive as the mussels supporting the horizontal spring have to be much bigger and stronger, but are also much more high performance and fast maneuvering oriented with the low profile and horizontal weight distribution of the long torso and long and flexible spine spring leveraging a large head and balanced by long tail.

The distribution of weight and the arrangement and length of the mechanical springs of the motor system in quadrupeds are correlated to the feeding strategy

Quadrupeds with vertically distributed rectangular profile (e.g. gazelle) form by long limbs and short and less flexible spine are energy conservation tactics for improving the locomotion EROEI when metabolizing products of lower energetic intensity (i.e. vegetation).

Horizontally distributed rectangular profile (e.g. leopard) of long and flexible spine and shorter limbs are designed for improving the feeding strategy EROEI from metabolism of higher energetic products (e.g. meat).

In most pronograde quadrupeds the head is proportional or relatively small and it is supported by horizontal neck and the muscles which hold it in position are dependent on the support of the shoulder muscles, working as a pulley system on the neck leveraged the spine and supported by the weight of the center and rear parts of the body, if the head continuously growing more mass must be installed and distributed to the rear or a longer torso and a tail must be added to leverage the added weight at the front to maintain the balance of the pronograde design usually for the purpose of maintaining agility with large head (as in Tigers).

When quadrupeds’ fitness strategy is based on big and heavy skull and growing in mass is not possible (due to phenotypic limitations, feeding strategy underlining, environmental, ecological and other factors) then another form of leveraging will developed based on leveraging the head weight with a pulley system shaped as an irregular quadrilateral shape where the rear limbs are shorter than the front ones and are vertically diagonal and the height difference is leveraged by longer spine in a slope from the front to the rear creating a triangular posture (as in giraffes to support the neck), the pulley is must be supported by weigh which achieved by unbalanced distribution of weight to the front of the body by enlarged muscular chest, shoulders and front limbs adding weigh to support the angular pulley of the spine all driving for centering the distribution of mass to the front part of the body toward above the front limbs.

An example of doing the process from rectangular quadruped to triangular we can find in the hyena with its big heavy head, that is not developed to contain large brain but for the purpose of mechanical bone-crashing pressure delivered by wide and massive jaw and supported by strong head and neck muscles. The hyena adapted their design for leveraging their large skull by short torsos and lower hind quarters, high withers and their backs slope noticeably downward toward their rumps, forelegs are high, while the hind legs are very short and their necks are thick and short creating triangular silhouette rather than rectangular as most quadruped.

But the hyenas, giraffes and other quadrupeds that moved their center of weight to the front and shaped their rear as a leverage in order to support heavy jaws or necks do not have the high energetic expenditure of supporting large computer in their skull as the apes does (i.e. chimpanzees) which put more pressure on the energetic input that is necessary to support it.

The limitations of the quadruped’s design on the maximal weight, shape and size of the head was also due to the physical limitation of the skeletal and muscular system to support increasing head weight and the maximal ability to compensate the head weight by moving more mass to the front for leveraging which eventually affect negatively the quadruped locomotion in terms of energetic efficiency, performance and even motoric functionality as a result of the growing imbalance between the front and rear part.

The process of transition from pronograde animal to orthograde (an erect or a vertical body position) animal present many physical and behavioral challenges to such transition that makes it almost unlikely that they can become fully orthograde or horizontally bipedal by transiting first to triangular posture where the back is slopping to the rear (were the spine makes the hypotenuse edge and front limbs are the opposite edge and the surface makes the adjacent edge) for that to happen an opposite process of moving the weight to the back and sloping from the rear to the front have to initially occur where the size of the head is getting smaller (e.g. bipedal dinosaurs with large heads like the t-rex started as small head animals that stand on their hind limbs to reach taller vegetation and eventually become bipedal and the spine of their quadruped early ancestors was sloping to the front and not to the back).

Our ancestors made the process of transitioning their locomotion from quadrupedalism to bipedalism directly as a phenotypic developmental strategy rather consequently because of their orthograde posture which started from their loris like ancestors locomotion strategy of clinging and climbing on vertical surfaces (tree trunks) as a result of extreme energy conservation strategy and later on maintain their orthograde posture as they transit to orthograde posture for arboreal brachiation of a gibbon like creature (due to their lack of tail which is crucial for the agile locomotion in arboreal quadrupeds such as lemurs) of perpendicular position of the spine where the neck is vertically balancing the rounded head as a ball on a poll.

The orthograde posture of our ancestors was an adaptation for arboreal brachiation and it was the main factor for the later adaptation for the terrestrial locomotion strategy of bipedalism driven by their existing perpendicular design and large head containing a large brain with processing power pushed to the extreme.

The large brain of our gibbon like ancestors was due to the high capacity of processing power that is needed for operating the mechanics of the motor system for performing extremely fast and agile brachiation in 3 dimensional environment of extreme complexity high above the ground in the rainforest’s canopy, where motile decisions are dependent on multiple streams of high volume raw data input from different sources are multithreading with multi-dimensional logic layers processing cognitive function fast enough to operate current gait while planning the rout and choreographing and synchronizing the next few steps of motile functions of point of contact for pendulum momentum and the projectile trajectories while traveling in speeds of more than 50 kilometers per hour.

Our ancestors didn’t grow big brains and skulls when they start walking on two rather they start walking on two as they were transitioning into an orthograde position suitable for ground walking utilizing their existing orthograde design adapted for arboreal brachiation (long hands, short legs, short torso) the triangular posture was imposed on them when knuckle-walking on the ground by their short hind limbs and long forelimbs and their large head was with a base centered to their spine rather than to the center of gravity consequently forming a triangular profile where the spine and neck are aligned and forming the hypotenuse edge sloping down to their hips in about 45 degrees.

The adaptation for bipedalism was more energetically cost effective and biomechanically plausible (maybe it was a cognitive strategy driven by the large size of brain to maintain itself?). Other factors also drove our ancestors to bipedalism such as the knuckle-walking position and angle of the neck and the pressure of the heavy skull on the weak neck muscles that become less developed as a result of the constant orthograde posture when hanging from branches, swinging and when sitting upright on top of branches always keeping the head and the neck center to the spine and gravity or to the center the centrifugal force (i.e. acts outward on a body moving around a center, arising from the body’s inertia) when preforming a pendulum movement.

 

Example of calculating the energetic expenditures of an animal behavioral activities during one daily cycle in order to calculate its Daily EROEI and Daily energy stock balance

In order to assess the evolutionary trends in a species of organisms we can factor all the physical properties, autonomic and rudimentary and biomechanical activities partaking in the total energy expenditure of the specific behaviors which are incorporated in the fitness strategy of the organism in the context of its environment in order to produce a profile which include the animal internal energetic consumption demand in correlation to it energetic balance and in correlation to it environmental energetic stocks and supply. The first stage is to establish baseline for the organism state at a certain life history stage at a certain point in time and space; assessing its short term EROEI at the end of its base metabolic cycle (can be daily as in primates, monthly as in some sharks, yearly as some reptiles etc.) here is an example of the methods and assumptions for a daily energetic expenditures of a fictitious tailless primate using brachiation as its primary locomotion strategy living in the equatorial broadleaf rain forest:

  • Mass units: using Kilograms (kg) on planet earth at sea level different then the newton weight (1 kg unit for a weight measure equals = into 9.81 N (newton earth)) although equivalent weight and mass unit type measure often used, the conversion for N is important for calculation of energy consumption when performing locomotion.
  • Temperature units: measured in Celsius for easy presentation but for calculation kelvin units will be used (0C=~273K, 25C=[25C]+273=~298K etc.)
  • Energy unit: using kilocalorie (kcal) as the unit of energy: the small calorie or gram calorie (cal) is the approximate amount of energy needed to raise the temperature of one gram of water by one degree Celsius at a pressure of one atmosphere. The large calorie or kilogram calorie (kcal) is defined in terms of the kilogram rather than the gram. It is equal to 1,000 small calories per 1 kilocalorie.
  • Daily cycle: a 24 hours cycle starts and ends at sunrise (e.g. in equatorial region the sun rise around 6:00AM: in that region the 24 hours cycle starts at 6:00AM and ends at 5:59AM the next morning).
  • Behavioral categories: foraging, eating, socializing, playing, resting, sleeping, reproduction etc.
  • Daily activities schedule (the time in the day and span of behavioral categories): includes the schedule and timespan invested in all the regular activities performed during one day and one night (such as foraging, eating, socializing, taking care of others, playing, resting, sleeping etc.)
  • Energetic baseline output: the sum of all energetic expenditure invested in autonomic bodily functions and behaviors that occur in a living organism (without conscious control) such as thermoregulation (the process of regulating the core temperature of the organism by generation, preservation and discharging of heat), metabolism for anabolic and catabolic activities that maintain the energetic balance of animals etc.
  • Behavior gross energetic output (per unit of time): the total amount of calories spent during a unit of time (one second/minute/hour) of activity and it is including the sum of all the energy invested in the systems that are actively participating in such activities (motoric systems, neurovascular systems etc.) and the energetic baseline output of all the autonomic and rudimentary bodily systems that are active during the behavioral activity, correlated and adjusted to a median temperature in the animal environment (e.g. 25C°, 35C°, 10C°, etc.), humidity and radiation intensity which are all part of the factors that are required in order to calculate the heating/cooling expenditures.
  • Daily gross energetic output (per behavior per day): The sum of the gross energy (kcal) invested in one category of behavioral l activity during 1 daily cycle
  • Total daily energetic output (per day for all activities): the sum of all the gross energetic expenditures (kcal) invested in of all behaviors performed during 1 daily cycle
  • Energetic input: the total amount of energy (Kcal) that was consumed digested and metabolized during 1 daily cycle.
  • Gross energetic stock: the total amount of energy (Kcal) that is stocked in the mass of an organism
  • Work energy stock: the amount of energy that is ready for the disposal by the organism for the purpose of executing and supporting behavioral and baseline activities
  • Daily EROEI value: the total amount of energy input in 1 daily cycle less the total daily energetic output during that cycle.
Setting energetic intensity scale and values per 1 hour of behavioral activity (adjusted to a median temperature in the animal environment of 25C°:
  • 1/Baseline/minimal intensity (Basal body temperature (BBT)): is the lowest body temperature attained during sleeping excluding initial digestive activities (50Kcal/hour)
  • 2/low intensity: in the example below I averaged all the intensity levels of activities involved in the behavioral categories of eating and socializing to 100Kcal/hour:
    • Rest/Stationary awakens state: (75Kcal/hour) measured immediately after awakening and before any physical activity has been undertaken excluding initial digestive activities.
    • Eating: (100Kcal/hour) food consumption and initial digestive activities during a mostly stationary state.
    • Slow locomotion: (125Kcal/hour) non-ricochetal brachiation (swinging) (or tailed primate walking on branches)
    • Low overheating/overcooling: ±2.5-5C° from median temp (125Kcal/hour)
  • 3/medium intensity: (200Kcal/hour) intensity level of activities involved in the behavioral category of foraging in the example bellow:
    • Cruising speed locomotion: low to medium speed of ricochetal brachiation (or tailed primate running on top of branches)
    • Medium overheating/overcooling: ±5-10C° from median temp
  • 4/high intensity: (300Kcal/hour) not applicable in the example bellow
    • Fast locomotion: medium to high speed of ricochetal brachiation (or tailed primate performing a series of sprinting for long distance leaps on top of branches)
    • High overheating/overcooling: ±10-20C° from median temp
  • 5/maximum intensity: (500Kcal/hour) not applicable in the example below
  • Fight/flight locomotion: include physical attacks and/or acrobatic high speed ricochetal brachiation when chasing or being chased (or tailed primate constantly sprinting, leaping and changing direction on top of branches in a series of short and long distance leaps)
  • Hypothermia/hyperthermia: over ±20C° from median temp
Calculating daily cycle energetic expenditures:

Observations
  • In order for the fictitious primate to maintain daily equalized EROEI it needs to consume 2,400Kcal/day in median temperature of 25C°.
  • The amount of energy invested in the consumption of 2,400Kcal daily is 1,400Kcal which includes the energetic expenditures of the behavioral activities of foraging trips and eating.
  • The amount of time invested in the consumption of 2,400Kcal daily is 8 hours which includes the span of the behavioral activities of foraging trips and eating.
  • The amount of energy invested in other daily behaviors is 400Kcal which includes the energetic expenditures of the behavioral activities of socializing.
  • The amount of time invested in other daily behaviors is 4 hours which includes the span of the behavioral activities of socializing.
  • The amount of energy invested in sleeping is 600Kcal which includes the energetic expenditures of the behavioral activities of socializing.
  • The amount of time invested in sleeping is 12 hours which includes the span of the behavioral activities of socializing.
Conclusions
  • Any decrease in the EROEI of the consumption activities due to change in the availability, distribution, range or the energetic value of the food sources due to change in abiotic factors, intraspecies, interspecies conditions etc., will increase the amount of time and energetic expenditure invested in consumption on behalf of the other daily cycle behavioral activities such as socializing and sleeping.
  • Any increase in the daily energetic expenditures due to change in temperatures or precipitation will increase the amount energetic expenditure invested in thermoregulation or in trips for maintaining dehydration will demand more consumption of energy daily and will result in an increase in the amount of time and energetic expenditure invested in consumption on behalf of all other daily cycle activities.
  • Any introduction of energetically expensive behavioral activities such as fight/flight demand an increase in the amount of time and energetic expenditure invested in consumption on behalf of the other daily behavioral activities such as socializing and sleeping (fight/flight activities can also result in injury which will affect the daily EROEI by demanding investment of energy and resting time for the recovery process)

Each change in the daily energetic balance and EROEI that will persist due to one, some or all of the above conclusion (as well as many other unmentioned factors that may affect the energetic balance in a negative or positive way), will affect the fitness of an animal and depending on the rate of escalation of the conditions that negatively affect the daily energetic balance and EROEI may ignite a change in the fitness strategy. A species can maximize its energetic equilibrium without physical changes but only for a short term by reducing its energy output in the form of time, in some behavioral activities and shifting it into activities involving in energetic consumption. There are other methods of increasing the energetic input and they are dependent on many colluding biological, cognitive and environmental factors which enabling or restricting behavioral adaption technics.

 

Aggression a key driver of evolutionary adaptation as a process of arm-race adaption (predator and prey) and as a responses to the introduction of threats into specie’s habitat

Changes in the energetic balance in a species of animal may suggest a beginning of an evolutionary adaptation process to regain energetic balance such balance can be affected negatively by the introduction of new condition.

Like any other counter-fitness factors that are introduced into a species environment the arrival of an attacker can ignite evolutionary adaptation in the attacked which will take a rout of behavioral or physiological adaptation or both routs each of them possess a spectrum of possible developmental morphological directions and many methods for responding to a newly arrived aggressor; (determine by the genotypic and phenotypic assets of the attacked species) in the short term the response must be carried by existing assets and in the long term by development of more specialized assets and depending on the phenotypic range and genotypic potential of the physical and cognitive assets and traits the immediate responses that are based on existing assets are representing the spectrum of potential immediate responses that the successful ones will ignite a developmental processes to amplified the successful strategy, the responses can be mostly relay on one of the two main categories or in between:

Physical response for new threat if successful can indicate the emergence of a morphological adaptation: utilizing existing phenotypic assets in the form of phenotypic arsenal of weapons or features that can potentially be used as weapons or defense (size, weight, strength of mussels, fangs, claws, thick skin, poison etc.) that can be immediately put use or can be easily modify within few generation of sexual selection.:

Cognitive response for new threat if successful can indicate the emergence of cognitive adaptation: utilizing existing and potential cognitive assets in the form of an arsenal of behavioral traits and tactics such as organizing in groups (social organization) outsmart aggressors etc.

There are different behavioral options for responding to a threat can be in the category of fight or flight, the two amygdala based responses and their derivatives (intimidation, hiding, migrating etc.). The developmental level of the attacked will determine if the immediate response which will be predominantly relay on the most developed asset either cognitive or physiological:

Fight – confronting threat with aggression

Individualistic based strategy for responding with fight for new threat will be mostly relay on physical traits: to be able to defend means to have the physical properties and arsenal of physical weapons (fangs, claws, large and strong mussels, thick skin, poison etc.) equal to the threat or fighting tactics that can compensate for inferior physicality all have to be fully developed and operational at the time when the new threat arrives or else to be able to develop or enhance existing phenotypic weapons before their species is decimated by the predator.
Basing the fight strategy on phenotypic physiological strategies is in the roots of solitary animals especially the ones that are of predatory nature or which their reproduction strategy force them into fierce fights, sometimes to death for the opportunity to reproduce or to confront predators on regular bases especially when parenting etc.
An adaptation process for developing phenotypic weapons for fight can also be triggered by the most common type of fight – the fight between male of the same species for the opportunity to breed with the specie’s females or to defend a territory with females and juvenile members of their species (dimorphism; where the males evolved to have larger size and weapons that makes their phenotypic physiology (as well as their behavioral traits) different then of the females’) or protecting a territory of resources, which are all depended on solitary male ability to survive by itself and fending from predators until it wins a vicious fight with another male of its own species for the opportunity to breed which many times come with a territory that it need to defend from contester males and with a duty of defending a group of females and offspring from predators.
Solitary animals are usually more in-tune with the arm race with their own kin or with existing prey or predators, and as they can only rely on themselves and with generation they have to evolve and develop technological advantage to counter their attacker or rival technology or they parish, limited by the availability of environmental energetic products and conditions and by their ability to maintain positive energetic balance and EROEI which is crucial for developmental evolution.

Group based strategy for responding with fight for new threat will be mostly relay on cognitive traits: means the members of the species are organized in groups prior to the arrival of the aggressor or have the cognitive capacity to form groups in order to gain the advantage of the combined mass, strength and stamina in order to attack or defend themselves.
There are different behavioral adaptations which are the premises on which aggression groups are formed for the purpose of defending or attacking and they are dependent on the premises of which aggression based groups are formed:

  • Hunting: the premise for attacking members of other species for the purpose of food consumption.
  • Cannibalism: the premise for attacking members of own species for the purpose of food consumption.
  • Defending own life: the premise for defending from members of your own species or other species for the purpose of surviving in the short term from becoming the prey of an attacker.
  • Defending resources: the premise for defending from members of your own species or other species for the purpose of surviving in the long term by not succumbing to energetically deficit due to the unavailability of the lost resources etc.
Flight – escaping confrontation:

Individualistic based response for flight will be mostly relay on physical traits: to be able to escape means to have certain phenotypic traits (light weight, long legs, strong muscles, wings, strong hind legs for jumping etc.) at the time when the new threat arrives, or to be able to develop or enhance existing traits before their whole species is decimated by the predator.

Other mostly individualistic strategies for dealing with aggression are avoiding confrontation by staying out of sight, out reach, or by being energetically worthless by adopting morphological or behavioral strategy which intended for lowering its energetic value, increase the energetic cost of acquisition and consumption resulting in negative EROEI for the attacker in other words the energy invested by a predator in acquiring and consuming the animal is higher than the energetic return from its prey. Animals can avoid being attacked by adopting various tactics and strategies which reduce the predator EROEI (by reduce mass, vast distribution, stamina etc.), can cause an energetic or physical mutilation to an attacker (by developing a defense system such as spikes, shell, poison etc.), that can increase the energetic expenditures of foraging and locating them by predator (such as living underground, camouflage, becoming nocturnal etc.) all of the needs a major phenotypic adaptations process to begin with and constant fine tuning of their strategies in accordance to their attackers development in the long run.

Group based response of flight will be mostly relay on physical traits: basing the survivability on the flight strategy which includes phenotypic physiological strategies is in the roots of group animals especially the ones that their physical properties are based on a less energetically efficient digestive systems due to the lower energetic values and the high energetic expenditures involved in the digestive process and metabolism of vegetation mater, or if their size and power are inferior to that of their predator and that their reproduction strategy do not include physical fights for mating (ritualistic fight).
In order to be able to effectively respond to a new threat the members of the species are organized in groups prior to the arrival of the aggressor or have the cognitive capacity to form groups in order to gain the advantages of the numbers.
There are different behavioral adaptation that are needed in order to form groups for the purpose of escaping attackers and they are dependent on the purpose and the premises of which such groups are formed which are utilizing or enjoying the advantages of some of the following tactics:

  • Dilution effect: animals living in a group “dilute” their risk of attack, each individual being just one of many, group provides benefits to the individual rather than to the group as a whole, which becomes more conspicuous as it becomes larger (e.g. the shoaling of fish).
  • Positioning: seeking a central position in a group to reduce the individual’s risk and to stay out of the area within the group in which the individual is more likely to be attacked by a predator. The center of the group has the lowest domain of danger, so animals will constantly strive to gain this position.
  • Improved vigilance: groups are able to detect predators sooner than solitary individuals. For many predators, success depends on surprise. If the prey is alerted early in an attack, they have an improved chance of escape.
  • Confusion: Individuals living in large groups may be safer from attack because the predator may be confused by the multiple individuals constantly changing positions, moving around and crossing the predator line of view, and as the group moves, the predator has greater difficulty targeting an individual prey animal.

Obviously more opportunistic species with wider genotypic range of phenotypic arsenal of physical traits, metabolic gamut and behavioral strategies have higher spectrum of abilities to possibly adapt to new conditions than more specialized spices that are centering their fitness strategy on relying on specific traits consequently narrowing their phenotype for specific ecological niche which ultimately leading to a bottleneck or a threshold from which adaptation becomes impossible.

Aggression in non-predatory species are a direct or indirect derivative of environmental condition and it is one of the main drivers of phenotypic evolution

Aggression is the most counter fitness factor as it represents an energetic burden (that is comparable for extreme abiotic condition and extreme recession of ecosystems) in the heavy toll it presents not only on the energetic balance and EROEI of the individuals of a species, but on the fundamental premises of survivability of staying alive and reproduce, aggression have to be taken in consideration in every adaptation which meant to improve the fitness, aggression is one of the main threats for fitness.

Aggression as one of the important evolutionary drivers

The development of fitness strategies are always in consideration with the premise of aggression as the determining factor in the evolution of all species.

Aggression influences all other fitness strategies some are derivatives of dealing with aggression: such as defending from aggression, hiding from aggression, avoiding aggression and escaping from aggression, while other are strategies of utilizing aggression for achieving energy and reproduction.

Aggression is either the secondary or the main driver for fitness strategies in all of the predatory non-predatory species

Aggression is executed for many reasons and there are different levels of aggression executed to achieve a certain functional and operational outcome:

Killing physically or energetically: for the purpose of resources acquisition that can be the flesh or physical resources in the possession or in the access of the attacked such as depriving them from food sources in a territory, food sources acquired or in the acquisition of the attacked and for reproduction opportunity (e.g. a lion killing leopard for its prey or taking over a pride for resources and reproduction)

Wounding/crippling physically or energetically: for the purpose of breaking up an attack (defense) or for the purpose of defeating in order to depriving the attacked from access to resources and reproduction opportunities (e.g. a lion chase the previous patriarch from its pride and territory)

Intimidating: utilizing passive aggressiveness by a display of power (e.g. individual mass, size, strength, or weaponry or by ganging in groups) or active aggression displays of preforming tactics of attack and restricted display of violence, or by limited acts of violence for the purpose of exampling individuals for creating systems of laws of punishments of individuals for purpose of depriving others from access to resources and reproduction opportunities or for forcing them to share the resources they possess with minimal resistance.

Forms of aggression based strategies

All the above forms of aggression may have different levels of short terms impact on the survivability of an animal (e.g. death, injury or banishment) but all levels of aggression are equalize in their impact of longer term on the survivability of individuals, groups and species due to the energetically toll inflicted on the attacked by the diminishing of their fitness (starving, death from injury, disruption of reproduction etc.) which ultimately jeopardize the longer term survivability of the species and reproduction possibility so all levels of aggression can all be considered as counter fitness in their ultimate results on the attacked.

There are two categories of group organized aggression (not considering the intraspecies aggression between individualistic males for the opportunity to become a single Alpha male and breed), between species and within a species and they represents two very different types of fitness strategies based on organization of individuals aggressive or defensive groups and for the group tactics and strategies they utilizing to achieve their purposes:

Interspecies aggression driven and driving fitness strategies:
  • Interspecies attack group: group that is organized for the purpose of attacking members of other species
  • Intraspecies defense group: group that is organized for the purpose of defending themselves against aggression executed by members of another species
Intraspecies aggression driven and driving fitness strategies:
  • Intraspecies attack group: group that is organized for the purpose of attacking members of its own species
  • Interspecies defense group: group that is organized for the purpose of defending themselves against aggression executed by members of another species

Most of the animals are organized internally in accordance to interspecies factors with the normal predator and prey arms race but a small number of species is utilizing to a certain extent, the fitness strategy that is based on intraspecies aggression and only two animals today are basing their social organization on intraspecies aggression as the driver for forming a group.

When answering aggression with aggression both the defender and the attacker have to utilize aggression as their methodology that may differ in tactic.

Fitness strategies to oppose predation which are based on confluence of the fight and the flight strategies with combined physical and behavioral tactics which are implemented dynamically in accordance to specific situations (i.e. fight when you can win or flight if you can’t), signaled the path for the evolution of higher levels of logic where optimized and adjusted sets of tactics are deployed in correlation and in accordance to each particular or circumstantial requirement in the most energetically efficient method.

From visceral motility to logical motility

Behavioral responses are determine by cognitive assets Evolution is a linear process when it comes to morphological changes and exponential development when it comes to cognitive development with laws limiting and governing the type of cognitive fitness strategies that are available for the specimens