A biotic regulation perspective on Nate Hagens’ Behavioral Thermodynamics: the Maximum Power Principle, and how life defends itself against its biggest consumers.
Anastassia, you are indeed correct, this is not a Christmas bauble. Instead a gem of much greater significance and brilliance. "A similar impairment occurs in the economy under monopolization, which in effect represents the same process: a transfer of power from a large number of small actors to a single large one."
I am going to borrow that graph that shows that roughly 90% of the primary energy is consumed by organisms 0.1mm or smaller for my Agricultural audience. (I assume that graph is for all life on earth, but that it still applies approximately to terrestrial life?) At least here in Southern Africa, we (humans) cleared our agricultural land from woodland that was photosynthesizing for perhaps 300 days a year. We now grow an annual summer crop which photosynthesizes for less than 100 days, but then we send at least half that energy into the towns and cities with our agricultural produce. So our soils have to survive on 1/6th of the primary energy they evolved to need. It is little wonder they are dying.
Although developed human societies will switch from fossil fuels to nuclear, the under-developed societies will continue to burn wood. There is already an area near me where the community has resorted to burning cow dung - which will accelerate the death of their soils even faster.
Your first graph (showing the band in which life exists, between about 0.7 and 80W/kg) is interesting enough but I propose that the lower limit down to the 0.1W/kg is simply because of the limited patience of human scientists to look that far. The upper limit? Possibly because of the challenges of disposing of the waste metabolic heat? The few outliers above the upper range, possibly exist in extreme environments like oceanic thermal vents.
Finally, for now, can you (or any of your audience) explain why there is no discernible recovery of the Blue Whales? I guess I could ask AI, but I would prefer a human answer.
Have a great 2026 everyone, Bruce Danckwerts, CHOMA, Zambia
Re Blue whales.. perhaps it has to do with the length of their breeding cycle.. and the fact that the foods they prefer are now being 'mined' commercially by humans. ^..^
Or their nutritional needs are being denied altogether with the inclusion of thousands of dams worldwide. There's enough evidence/ proof that in the subarctic/Polar regions most of the freshwaters for the entire Northern Hemisphere has been impounded behind 17 major hydroelectric dam installations, practically surrounding the Arctic Ocean and this has been occurring now for almost 60 years
Commercial fishing wreaks havoc and absolute destruction on the oceans and marine life especially bycatch and ghost fishing. We need to change our ways while we have time. We need to choose more plant based and cultivated foods that are sustainable and just for all.
I quit commercial fishing in the ‘70s because of what I saw.. the overfishing, the ‘bycatch’ (and its disposal processes), the effects that logging had on streamflows where I lived (next to salmon-bearing rivers & creeks), other changes.. prices, weather, attitudes, a ‘feeling’ among many of us that “something is really wrong” with our ‘industry’. I was also a tree planter in those days.. & was watching the effects that logging had on spawning streams, ‘up close & personal’. ^..^
Bruce, the graph shows what can be called "normal" metabolic rates, also referred to as basal or endogenous rates. These are measured for organisms at rest and some time after feeding, since eating temporarily raises metabolism.
You are right that there are both lower and higher extremes, and we did spend time looking at those as well. In particular, there are two regimes of very low metabolic rates. One is seen in animals like hibernating bears, which keep energy consumption extremely low while still maintaining order in their tissues. The second is an almost complete shutdown, as in aestivating snails or dormant spores. In this state, damage slowly accumulates, but energy use is essentially negligible, on the order of a thousandth of a watt per kilogram or less.
For details, see Makarieva A.M., Gorshkov V.G., Li B.-L., and Chown S.L. (2006), Size- and temperature-independence of minimum life-supporting metabolic rates, Functional Ecology, 20, 83–96: https://bioticregulation.ru/ab.php?id=fe06
At the other end of the spectrum, the highest metabolic rates occur in actively growing bacteria and in the tissues of flying animals, where power consumption can exceed a thousand watts per kilogram, see
Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Energetics of the smallest: Do bacteria breathe at the same rate as whales? Proceedings of the Royal Society of London, Biological Series, 272, 2219-2224. https://bioticregulation.ru/ab.php?id=bact
Interesting. The other interesting feature of that first graph, is the backward slope of the blue (Endotherms - I assume that means warm-blooded?) Is there an explanation as to why the smaller Endotherms have a higher metabolic rate - just the scale effect that smaller animals have more surface/kg than big animals and so lose heat quicker, requiring a high metabolic rate? There seems to be a (steeper) slope to the Insects and Copepods (but that could be because their data is hidden behind the Insects, with whom they overlap). The prokaryotes slope just slightly the other way. None of this changes your main article, just interesting to note the differences.
There is a HUGE and at times exceedingly sophisticated literature devoted to deciphering the patterns by which metabolic rate scales within different taxa. This work focuses not on absolute metabolic rates, but on the scaling exponents—that is, how rapidly metabolic rate changes with body size.
Basically, you are right: it all comes down to the surface-to-volume ratio. In warm-blooded animals that maintain a more or less uniform body temperature, heat is released at an approximately constant rate per unit surface area. As animals become smaller, they therefore must produce more heat per unit mass, exactly as you say.
For cold-blooded animals, the same logic can be applied to oxygen consumption, which is also surface-specific and ultimately governed by diffusion. As an organism becomes larger, it has a lower oxygen supply per unit mass and must therefore metabolize more slowly. If environmental oxygen levels increase, it follows—given a universal metabolic constraint—that larger organisms can grow even bigger without leaving the optimal range. This likely explains why ancient dragonflies could reach wingspans exceeding one meter.
Likewise, an increase in ambient temperature can also extend maximum size. We had another publication on this topic re maximum sizes -- Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Temperature-associated upper limits to body size in terrestrial poikilotherms. OIKOS, 111, 425-436. https://bioticregulation.ru/ab.php?id=wwloi
I remember watching a talk from Dana Meadows (Limits to Growth) regarding systems thinking in which she discusses fish stock collapse. Often the signs that we are overfishing come far too late to allow their populations to recover.
(the lecture is well worth watching, if you search Dana Meadows Sustainable Systems - it's the one "Presented at the University of Michigan Ross School of Business" in four parts)
It's a fantastic book by Joe Roman, sorry the full title is "Eat, Poop, Die: How Animals Make Our World".
It describes how important animals are to nutrient cycling, for instance how nutrients get to high mountain tops via birds, how bears eating salmon bring nutrients to the trees, how whales move nutrients through the oceans, feeding in colder waters and then breeding (and shitting lots) in warmer areas, etc etc.
My favourite is when he talks about the midgies in Greenland ( I think) and how they will actually contribute to really healthy grass because they bring up nutrients from ponds and then shit and die all over the grass.
I think of it as "stirring the pot" - migrating animals could (over millennia) be a very effective way of moving essential nutrients from areas where a particular nutrient is abundant, to another area where it is scarce. Two problems in this modern world: Too many animals are now confined to game parks or reserves and so can no longer migrate. Also, one of the biggest sources of excrement (city sewage) is not recycled onto land, but flushed out to sea. When we do attempt to recycle it, we find it is contaminated with heavy metals (and a whole host of other unnatural chemicals - like hormones and detergents) that make them less suitable as fertilizer. Mother Nature does her best to glean these nutrients from the city sewage, and not only fish, but any seafood that humans bring ashore, bring these nutrients back onto land . . . . . but I live about 1,500kms from the ocean, and very little seafood makes it to my farm, let alone my closest town. And no fish make it. So, although I think animals are essential to any farming system, we still have to be prepared to address nutrient deficiencies that our animals, birds and insects have been unable to resolve.
I did some engineering work in the UK wastewater industry for a small time and I found it absolutely fascinating (though site visits would leave the stench reminder in my nostrils for days), everything was biological, leaving microbes and sunlight to clean the water. The resulting clear water flowed back into the river and I believe the solids was used by farming to fertilise their fields (not without the contamination issues you mentioned!).
One of the first things I did when we moved to our land here in the Philippines was to begin a composting toilet. It's such an underutilised resource - a great book explaining it all is Joseph Jenkins' Humanure Handbook which is available free on the internet. My composting methods are of course different as I believe he was in a temperate climate, but here in the tropics I utilise rhino beatle larvae to finish all my composts, whether it be humanure or the cow poop.
Another book that I digested many years ago was Steve Solomon's The Intelligent Gardener, he was all for re-mineralising the soil using rock dusts, etc. His context was a garden, not a farm though. I never ended up doing any of this since getting a soil test with the details needed was out of the question here (at the time, and probably still) and obtaining the necessary minerals was also pretty much out of the question.
I gave all the remineralisation ideas up after the local volcano, 30km away, blew up and gave us all a nice remineralisation. We were lucky, and only got a light dusting of ash.
Then I started listening to soil scientists like Elaine Ingham and Dr Christine Jones and started to get quite confused as I think Elaine's premise was that all the necessary minerals are always in the soil, they just need unlocking.
These days I don't know what to believe or follow but it doesn't stop me from following just basic soil principles, living mulches (my cover crops are weeds haha), keeping the soil always covered, integrating animals, minimal disturbance, living roots, etc etc. I wish at the start I'd had the mindset to record what the conditions were initially like (I had to leave a hose on overnight soaking the ground where I was digging my first veggie beds, it was that hard!). These days we have much better soil and our flooding is reduced. I'd be scared to take a comprehensive soil test though, haha.
We could be accused of drifting off topic ;-) but in fact it is Life and all of Life is related! I too am skeptical about Elaine and Christine's claims. Elaine is on record as claiming there are not 13 soil nutrients but 32 (or is it 33?) but I cannot find her list, nor any description of what function these extra nutrients do in the soil. It is statistically VERY unlikely that every soil has every nutrient available in roughly the right ratios. (Christine did admit to me that there are regions of the World known to have shortages - like Southern Africa is short of Boron and Selenium . . . and others.) Christine lost my following when she claimed that sea water was good for pastures! (a) as I mentioned, I am 1500km from the sea, and (b) although an application of sea-water might indeed raise the Brix levels of a pasture, it is probably the poor plants trying desperately to compensate for the near fatal dose of salt. I cannot remember the author, but I think the book was called Growing Life, in which he gave his minimum nutrient levels for what he regarded as a fertile soil, on which one might grow organic food. My K was 62% of his minimum, and my Ca, P, S etc were less than 10% :-( Because we send nutrients to the cities with our produce and most cities do not send them back, I do believe on my soils I HAVE to be alert to the possibility of nutrient deficiencies. You can read my approach to soil management on the Farmers Handbook page of www.radio4pasa.com - look for the article, Farming as a System. Another one discusses nutrients, and will give you the average levels on my farm - leached, ancient, granite sands.
I think hopefully Anastassia will agree we are very much on topic to be talking about caring for soil!
Bruce, loved your Farming as a System document, everything I agreed with and had also come to the same conclusions from all my reading. Wish I'd read your document back when I was starting... to have it all in one place is so valuable.
We're in a very different context but the principles are the same regardless. I also love how you put in the social and cultural aspects too, so important.
Jeff Lowenfels did some great books; Teaming with microbes, Teaming with nutrients, bacteria, etc which you might find useful regarding the soil biology. Think I only read the microbes one, I should go back and read the rest.
Re the quote that B. Danckwerts commented upon, "..a similar impairment.." etc would appear to hold true for MONEY, these days.. as a very few billionaires are growing wealthier at an exponential rate, as the middle class is reduced, while the poor gain little or nothing. ^..^
Took the words right our of my mouth John. It's becoming increasing obvious that fewer people are gaming the economic system, getting wealthier.
The economic model of our day is extraction. A system that ultimately throws the whole planet, our species, and all life under the bus.
Extraction without replacement, the planets thermodynamic requirements are being overridden by an economy of extraction. Will more man-made blunders appear again or will we work within the planets thermodynamic requirements for resilienceand continued health?
Leon everything is converging! I'm a climate researcher my associate and friend also is a climate researcher however his expertise is whole system integration and analysis. He too has a substack...Regeneration. He could not help but see a significant pattern emerging with our economy and with Trumps assistance helping to rush us over a deep cliff
It might also be when you read widely enough you can see commonalities in everything!
I'll try to have a read of the link you sent, it's a long one. I've been following a great substack called System Failure by Nathan Knopp and it's basically covered history from why the Roman Empire collapsed, the rise of Christianity as the new Empire, the collapse of their authority after the black death plague, the rise of Science as the new Empire after enlightenment, etc etc. Nathan only writes a short 5 - 10 minute read on each essay so easy to digest and get through, it's very interesting and makes what's happening now more understandable, only this time civilisation is globe-spanning.
Its also possible to skim through the most important points of this. He actually just released a followup article as well. I think that the followup lends itself easier to skim most of the salient points and he too refferences histories similarities as empires rise and fall.
Thank you, I also like this connection. From my perspective, enshittification is strongly linked to a declining surface-to-volume ratio. Consumers are at the surface, and they receive progressively less attention and poorer services. The same applies to large animals: the environment becomes something external, and they are no longer able to regulate it.
This is an incredible set of active models allowing reflection on the larger systems, very thought provoking. Looking forward to reading more as further development of the model(s) you are connecting this readership to appears here in print.
How do we know that the shared metabolic rate isn't simply the convergent property of organisms jamming themselves as close as they can get to the ceiling for dissipating waste heat before their proteins lyse without going over?
Proteins start to break down at fairly high temperatures (around 40 °C), but all the organisms shown in the first figure live well below that range (most are measured at 25 deg C) and yet display similar metabolic rates. That makes it unlikely that the shared metabolic rate simply reflects a hard thermal failure limit.
Instead, it looks like the largest animals had to become endothermic to compensate for the size-related drop in mass-specific metabolic rate by generating extra heat and to catch up with the rest of the biosphere. Without doing that, they would fall outside the optimal metabolic range and would likely be at a competitive disadvantage. This is, by the way, a strong argument in favor of dinosaurs being endothermic, something that used to be considered as a heresy but gradually accepted.
Anastassia, many thanks for yet another thought-provoking article; it kept my mind busy since its publication. A couple of thoughts.
First, the consumed shared productivity over the characteristic length of a body is some kind of x^-k pattern (k being some real number). This makes sense to me for a constant input power through the sun (perhaps of the order of 200-250W/m^2 at the ocean surface) and conversion of that energy to biomass by the smallest entities, e.g. phytoplankton, at about 0.5-1W/m^2. Only a fraction of that generated stock can sustainably be ingested by larger bodies at a certain conversion rate or efficiency. That said, having a tally of all organism in size bins will allow to determine k, something > 0. However, Helen Chersky posits in her book Blue Machine that the total biomass is constant across different sizes. With a constant relationship between body size and energy consumption, k would then be rather 0; every size category would need to consume the entirety of the next smaller category, which does not sound plausible (unless it happens once). What are your thoughts on that?
Second, one of your graph shows that humanity consumes about 10% of the energy share where a more appropriate sustainable share would be sub 1%. It follows that the human population is about one to two orders of magnitude to large, or will be once the fossil energy bonanza is finished.
Thank you, Peter, for these thoughtful comments. The claim of constant total biomass is usually based on estimates of a scaling exponent that is statistically indistinguishable from zero (or -1), obtained from a linear regression of log-transformed biomass (or population density) against log-transformed body mass.
For consumed productivity, a decline of about two orders of magnitude over at least four orders of magnitude in linear size—and twelve orders of magnitude in body mass—would imply a biomass–body-mass scaling exponent of −0.17 (or, for population density against body mass, -1.17) across the full range. Because this range is rarely considered in its entirety, such a trend can easily be obscured by limited or noisy data. Thus instead of extrapolating from allometric analysis, a more robust method is a direct calculation of energy consumption for specific classes of consumers, as I did in the post and we did in our publications.
Furthermore, as we discussed in this publication https://bioticregulation.ru/common/pdf/ec04.pdf , there is evidence that biomass distributions are steeper in more stable ecosystems than in less stable ones, a factor that is almost universally ignored.
Anastassia, thanks for your detailed response; I'll study the paper and other literature, considering the likely relationship of body mass is probably proportional to body size to the power of three, or thereabout.
As for the second part, the consequence of a sustainable human population of about say two orders of magnitude less than would also mean that even a transition along a logistics curve towards stabilised population is unlikely (SW in LtG, 1972); rather a contraction to that stabilised world is to be expected.
Thank you for this excellent and thought-provoking writing about energy consumption. I need a little more explanation about your comment near the end of your piece: "Trees are not a resource." Since you stated what they are not, I really hoped your next statement would succinctly say what they are. This will help readers like me more easily share some concrete examples of what trees are. For example, are they biotic regulators? It seems, from your research, that trees are central to the biotic pump concept, particularly when they are part of an intact forest ecosystem. Does your statement that they are 'not a resource' argue for thinking of trees as much more than something to be exploited? Or, am I misunderstanding a major point in your writing?
Thank you for this feedback—much appreciated. Yes, you are right: trees, when they are part of a natural ecosystem, are major biotic regulators and an integral component of the biotic pump. They are, quite literally, the basis of life on land.
At the same time, I feel that these words are still too weak to fully express what I mean. That is why I deliberately left the phrase “trees are not a resource” standing on its own. My hope was that, rather than offering a ready-made conclusion, it would prompt readers to pause and think for themselves—something like opening a gestalt.
Sometimes, when a picture is complete, it appears and disappears just as quickly. But when something feels unresolved, it can linger and quietly irritate the mind for much longer. With my writing, I want to encourage people to think about the natural world in new ways.
Maybe nothing is a Resource. In 19th century classical industrial capitalist economics in which our society is still largely based, this term means "of exploitable nature" (rather than the previous meaning of an "inner strength"; Hausmann, Perreaux 2020), and to "allocate scarce means to *satisfy unlimited wants*" . This sounds like a delusional worldview resulting, perhaps, from a misplaced perception over our role on the planet. The world does not belong to us ! Life does not belong to us ! We can use what we need within balance with every other planetary inhabitant, and we are now in a better position to understand our limits to growth in both an economical and ecological sense. Unlike other species, we were able to overextend our population but now more able to keep it in check through awareness and nature-wise decisions.
Thank you for this. I'm reminded of a 2007 study in PNAS that found Human Appropriation of Net Primary Productivity is 83% in agricultural areas, and about 28% globally (that's a terrestrial fraction, I assume). That deeply brought home to scale of the ecological crisis to me.
There are also areas from which we do not consume primary productivity, but which we have anyway destroyed. This is a separate count. E.g. if you cut down trees for development, then there are no more trees which you could cut, so on this area you won't consume anything. Yet it will be destroyed and not contributing to climate stability. So there are unfortunately multiple metrics.
If you haven’t already, you should read some of Ernst Götsch’s writing on this subject. In his view, animals appear in contexts where the local net primary production has exceeded local metabolic capacity and the size of the animals is proportional to that excess. This allows for living systems to continually increase their production over evolutionary time and create larger stocks of “natural capital”. Of course these large movers of energy need predators to manage them and ensure they are working, in service as it were, to the greater system. Far from destroying vegetation, well managed large herbivores can dramatically increase NPP through a variety of mechanisms.
I would also add that large animals play a critical role in the creating more land. When large numbers of large animals are living, continental erosion rates increase (this happened with dinosaurs and Pleistocene mammals), but not above soil creation rates. Because of isostacy this means that the continent actually expands and more fertile land is created where plants can grow (erosion creates fertile deltas and fluvial depositions etc).
Interesting. Makes me remember a visit to farmers in Cameroon Highlands in Malaysia. I was horrified by the level of erosion by their farming of steep slopes. But one farmer explained to me that they got a much bigger and fertile valley bottom as a result. Also in Japan I remember similar expressions by farmers: "we cultivated away a hill"....
dear Anastassia, thanks for your contributions. I hope you keep going! In a paper from 2014 Haberl,Helmut, Karl-Heinz Erb och Fridolin Krausmann Human Appropriation of Net Primary Production: Patterns, Trends, and Planetary Boundaries, Annual Review of Environment and Resources Volume 39, 2014. estimated what they define as Human Appropriation of Net Primary Production (HANNP) to be 25 percent by 2005. Are you familiar with their calculation and if so, can you explain the difference between their estimate and the estimate by Viktor Gorshkov, apart from the fact that they are about the situation at a different times?
Thank you very much, Gunnar. I also hope to keep going, even if it has been a major shift in activities for me which I am still learning how to handle.
To compare Table 1 for HANPP from Haberl et al. 2014 with Gorshkov's estimates [https://bioticregulation.ru/common/pdf/ufn80-en.pdf, Section 11D ] it is useful to keep in mind that 1 PgC/year = 1 GtC/year = 10^12 Watt = 1 TW.
Gorshkov divided HANPP in two parts, one is power consumption due to human food, cattle fodder and timber, around 5 TW or 5 PgC/year, or 8% of terrestrial NPP (which he estimated at 60 PgC/year). This "steady-state" consumption is what is shown in the graph, to compare it with steady-state consumption in natural terrestrial ecosystems. This shows that there is an anthropogenic disturbance to stable architecture of energy flows even if the currrent food and sylviculture systems were "sustainable" in the short term.
The other part of HANPP is the decline in the biomass of the biosphere due to human disturbance (deforestation), which he estimated at 16 TW, or 27% of terrestrial NPP (giving a total of 35% for 60 GtC/year terrestrial NPP).
Vitousek et al. on which Haberl et al. are based considered not actual NPP but some potential NPP (about 75 GtC/year).
One more question: It seems as if your diagram on metabolic rate is contrary to the "mouse to elephant" curve as developed by Kleiber and others. Or do I misunderstand you or them?
The "mouse to elephant" curve is confined to the body size interval between the mouse and the elephant, and you can actually see it in the graph ("endotherms", also includes birds with a roughly similar scaling).
What matters is that this curve does not extend to all life, but breaks as one goes from one clade to another, and breaks in such a manner that every taxon approximately remains within the same interval of mass-specific rates.
Very interesting analyses and observations. However, as far as I can analyse, Max power still applies , following the original definition by Lotka: “evolution in a system aims to increase the total (bio) mass in a system, as well as to increase the flow of mass through that system, as well as the energy flux by increasing the system, as long as there is an unused share of mass and energy in that system. ”
And growth of power ends ( but not the principle) when equilibrium is achieved, which is a result of trial and error, growth and decrease of species follows availability and dependence of each other .
Humans finding fossil stocks and ways to use these disturbed the situation and fueled max power again. Until running out of fuel and collapse.
(By the way: and humans as well shaped a imaginary system of value by money , precisely to maximize power over energy and resources. )
This is someting that ancient Taoist master Zhuang Zhou (aka. Zhuangzi, Chuang Tzu) discussed 2300 years ago in a famous story about "Man who despised machine". When I first red the story it completely changed my life. In that story Zhuang Zhou argues that in using machines as prosthetics for resource overconsumption humans selfdestruct. What impressed me is the way that Zhuang Zhou relates machine use with morality. Wise man in the story explicitly said that machine use (even simple machines) causes overexpoitation of resources. That is a kind of cheating of Nature. He knows that work is easier done with machines but he doesn't want to use them because it is immoral. Taoists know that resource overexploitation is disturbing the energy flows of nature and that means big trouble for the menkind. Overconsumption brings arrogance and arrogance brings selfdestruction. Unfortunately, when you have competitor that overexploits nature (USA) and that is arrogant and aggressive, you don't have a choice but to follow the same path in order to survive. Your only chance is to build even more powerfull machines.
Anastassia, you are indeed correct, this is not a Christmas bauble. Instead a gem of much greater significance and brilliance. "A similar impairment occurs in the economy under monopolization, which in effect represents the same process: a transfer of power from a large number of small actors to a single large one."
I am going to borrow that graph that shows that roughly 90% of the primary energy is consumed by organisms 0.1mm or smaller for my Agricultural audience. (I assume that graph is for all life on earth, but that it still applies approximately to terrestrial life?) At least here in Southern Africa, we (humans) cleared our agricultural land from woodland that was photosynthesizing for perhaps 300 days a year. We now grow an annual summer crop which photosynthesizes for less than 100 days, but then we send at least half that energy into the towns and cities with our agricultural produce. So our soils have to survive on 1/6th of the primary energy they evolved to need. It is little wonder they are dying.
Although developed human societies will switch from fossil fuels to nuclear, the under-developed societies will continue to burn wood. There is already an area near me where the community has resorted to burning cow dung - which will accelerate the death of their soils even faster.
Your first graph (showing the band in which life exists, between about 0.7 and 80W/kg) is interesting enough but I propose that the lower limit down to the 0.1W/kg is simply because of the limited patience of human scientists to look that far. The upper limit? Possibly because of the challenges of disposing of the waste metabolic heat? The few outliers above the upper range, possibly exist in extreme environments like oceanic thermal vents.
Finally, for now, can you (or any of your audience) explain why there is no discernible recovery of the Blue Whales? I guess I could ask AI, but I would prefer a human answer.
Have a great 2026 everyone, Bruce Danckwerts, CHOMA, Zambia
Re Blue whales.. perhaps it has to do with the length of their breeding cycle.. and the fact that the foods they prefer are now being 'mined' commercially by humans. ^..^
Or their nutritional needs are being denied altogether with the inclusion of thousands of dams worldwide. There's enough evidence/ proof that in the subarctic/Polar regions most of the freshwaters for the entire Northern Hemisphere has been impounded behind 17 major hydroelectric dam installations, practically surrounding the Arctic Ocean and this has been occurring now for almost 60 years
Commercial fishing wreaks havoc and absolute destruction on the oceans and marine life especially bycatch and ghost fishing. We need to change our ways while we have time. We need to choose more plant based and cultivated foods that are sustainable and just for all.
I quit commercial fishing in the ‘70s because of what I saw.. the overfishing, the ‘bycatch’ (and its disposal processes), the effects that logging had on streamflows where I lived (next to salmon-bearing rivers & creeks), other changes.. prices, weather, attitudes, a ‘feeling’ among many of us that “something is really wrong” with our ‘industry’. I was also a tree planter in those days.. & was watching the effects that logging had on spawning streams, ‘up close & personal’. ^..^
Bruce, the graph shows what can be called "normal" metabolic rates, also referred to as basal or endogenous rates. These are measured for organisms at rest and some time after feeding, since eating temporarily raises metabolism.
You are right that there are both lower and higher extremes, and we did spend time looking at those as well. In particular, there are two regimes of very low metabolic rates. One is seen in animals like hibernating bears, which keep energy consumption extremely low while still maintaining order in their tissues. The second is an almost complete shutdown, as in aestivating snails or dormant spores. In this state, damage slowly accumulates, but energy use is essentially negligible, on the order of a thousandth of a watt per kilogram or less.
For details, see Makarieva A.M., Gorshkov V.G., Li B.-L., and Chown S.L. (2006), Size- and temperature-independence of minimum life-supporting metabolic rates, Functional Ecology, 20, 83–96: https://bioticregulation.ru/ab.php?id=fe06
At the other end of the spectrum, the highest metabolic rates occur in actively growing bacteria and in the tissues of flying animals, where power consumption can exceed a thousand watts per kilogram, see
Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Energetics of the smallest: Do bacteria breathe at the same rate as whales? Proceedings of the Royal Society of London, Biological Series, 272, 2219-2224. https://bioticregulation.ru/ab.php?id=bact
Interesting. The other interesting feature of that first graph, is the backward slope of the blue (Endotherms - I assume that means warm-blooded?) Is there an explanation as to why the smaller Endotherms have a higher metabolic rate - just the scale effect that smaller animals have more surface/kg than big animals and so lose heat quicker, requiring a high metabolic rate? There seems to be a (steeper) slope to the Insects and Copepods (but that could be because their data is hidden behind the Insects, with whom they overlap). The prokaryotes slope just slightly the other way. None of this changes your main article, just interesting to note the differences.
There is a HUGE and at times exceedingly sophisticated literature devoted to deciphering the patterns by which metabolic rate scales within different taxa. This work focuses not on absolute metabolic rates, but on the scaling exponents—that is, how rapidly metabolic rate changes with body size.
Basically, you are right: it all comes down to the surface-to-volume ratio. In warm-blooded animals that maintain a more or less uniform body temperature, heat is released at an approximately constant rate per unit surface area. As animals become smaller, they therefore must produce more heat per unit mass, exactly as you say.
For cold-blooded animals, the same logic can be applied to oxygen consumption, which is also surface-specific and ultimately governed by diffusion. As an organism becomes larger, it has a lower oxygen supply per unit mass and must therefore metabolize more slowly. If environmental oxygen levels increase, it follows—given a universal metabolic constraint—that larger organisms can grow even bigger without leaving the optimal range. This likely explains why ancient dragonflies could reach wingspans exceeding one meter.
Likewise, an increase in ambient temperature can also extend maximum size. We had another publication on this topic re maximum sizes -- Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Temperature-associated upper limits to body size in terrestrial poikilotherms. OIKOS, 111, 425-436. https://bioticregulation.ru/ab.php?id=wwloi
I remember watching a talk from Dana Meadows (Limits to Growth) regarding systems thinking in which she discusses fish stock collapse. Often the signs that we are overfishing come far too late to allow their populations to recover.
(the lecture is well worth watching, if you search Dana Meadows Sustainable Systems - it's the one "Presented at the University of Michigan Ross School of Business" in four parts)
Thank you, I will indeed look that up
Eat shit and Die Bruce
It's a fantastic book by Joe Roman, sorry the full title is "Eat, Poop, Die: How Animals Make Our World".
It describes how important animals are to nutrient cycling, for instance how nutrients get to high mountain tops via birds, how bears eating salmon bring nutrients to the trees, how whales move nutrients through the oceans, feeding in colder waters and then breeding (and shitting lots) in warmer areas, etc etc.
My favourite is when he talks about the midgies in Greenland ( I think) and how they will actually contribute to really healthy grass because they bring up nutrients from ponds and then shit and die all over the grass.
Please excuse my language.
I think of it as "stirring the pot" - migrating animals could (over millennia) be a very effective way of moving essential nutrients from areas where a particular nutrient is abundant, to another area where it is scarce. Two problems in this modern world: Too many animals are now confined to game parks or reserves and so can no longer migrate. Also, one of the biggest sources of excrement (city sewage) is not recycled onto land, but flushed out to sea. When we do attempt to recycle it, we find it is contaminated with heavy metals (and a whole host of other unnatural chemicals - like hormones and detergents) that make them less suitable as fertilizer. Mother Nature does her best to glean these nutrients from the city sewage, and not only fish, but any seafood that humans bring ashore, bring these nutrients back onto land . . . . . but I live about 1,500kms from the ocean, and very little seafood makes it to my farm, let alone my closest town. And no fish make it. So, although I think animals are essential to any farming system, we still have to be prepared to address nutrient deficiencies that our animals, birds and insects have been unable to resolve.
Absolutely agree.
I did some engineering work in the UK wastewater industry for a small time and I found it absolutely fascinating (though site visits would leave the stench reminder in my nostrils for days), everything was biological, leaving microbes and sunlight to clean the water. The resulting clear water flowed back into the river and I believe the solids was used by farming to fertilise their fields (not without the contamination issues you mentioned!).
One of the first things I did when we moved to our land here in the Philippines was to begin a composting toilet. It's such an underutilised resource - a great book explaining it all is Joseph Jenkins' Humanure Handbook which is available free on the internet. My composting methods are of course different as I believe he was in a temperate climate, but here in the tropics I utilise rhino beatle larvae to finish all my composts, whether it be humanure or the cow poop.
Another book that I digested many years ago was Steve Solomon's The Intelligent Gardener, he was all for re-mineralising the soil using rock dusts, etc. His context was a garden, not a farm though. I never ended up doing any of this since getting a soil test with the details needed was out of the question here (at the time, and probably still) and obtaining the necessary minerals was also pretty much out of the question.
I gave all the remineralisation ideas up after the local volcano, 30km away, blew up and gave us all a nice remineralisation. We were lucky, and only got a light dusting of ash.
Then I started listening to soil scientists like Elaine Ingham and Dr Christine Jones and started to get quite confused as I think Elaine's premise was that all the necessary minerals are always in the soil, they just need unlocking.
These days I don't know what to believe or follow but it doesn't stop me from following just basic soil principles, living mulches (my cover crops are weeds haha), keeping the soil always covered, integrating animals, minimal disturbance, living roots, etc etc. I wish at the start I'd had the mindset to record what the conditions were initially like (I had to leave a hose on overnight soaking the ground where I was digging my first veggie beds, it was that hard!). These days we have much better soil and our flooding is reduced. I'd be scared to take a comprehensive soil test though, haha.
We could be accused of drifting off topic ;-) but in fact it is Life and all of Life is related! I too am skeptical about Elaine and Christine's claims. Elaine is on record as claiming there are not 13 soil nutrients but 32 (or is it 33?) but I cannot find her list, nor any description of what function these extra nutrients do in the soil. It is statistically VERY unlikely that every soil has every nutrient available in roughly the right ratios. (Christine did admit to me that there are regions of the World known to have shortages - like Southern Africa is short of Boron and Selenium . . . and others.) Christine lost my following when she claimed that sea water was good for pastures! (a) as I mentioned, I am 1500km from the sea, and (b) although an application of sea-water might indeed raise the Brix levels of a pasture, it is probably the poor plants trying desperately to compensate for the near fatal dose of salt. I cannot remember the author, but I think the book was called Growing Life, in which he gave his minimum nutrient levels for what he regarded as a fertile soil, on which one might grow organic food. My K was 62% of his minimum, and my Ca, P, S etc were less than 10% :-( Because we send nutrients to the cities with our produce and most cities do not send them back, I do believe on my soils I HAVE to be alert to the possibility of nutrient deficiencies. You can read my approach to soil management on the Farmers Handbook page of www.radio4pasa.com - look for the article, Farming as a System. Another one discusses nutrients, and will give you the average levels on my farm - leached, ancient, granite sands.
I think hopefully Anastassia will agree we are very much on topic to be talking about caring for soil!
Bruce, loved your Farming as a System document, everything I agreed with and had also come to the same conclusions from all my reading. Wish I'd read your document back when I was starting... to have it all in one place is so valuable.
We're in a very different context but the principles are the same regardless. I also love how you put in the social and cultural aspects too, so important.
Jeff Lowenfels did some great books; Teaming with microbes, Teaming with nutrients, bacteria, etc which you might find useful regarding the soil biology. Think I only read the microbes one, I should go back and read the rest.
Re the quote that B. Danckwerts commented upon, "..a similar impairment.." etc would appear to hold true for MONEY, these days.. as a very few billionaires are growing wealthier at an exponential rate, as the middle class is reduced, while the poor gain little or nothing. ^..^
Took the words right our of my mouth John. It's becoming increasing obvious that fewer people are gaming the economic system, getting wealthier.
The economic model of our day is extraction. A system that ultimately throws the whole planet, our species, and all life under the bus.
Extraction without replacement, the planets thermodynamic requirements are being overridden by an economy of extraction. Will more man-made blunders appear again or will we work within the planets thermodynamic requirements for resilienceand continued health?
I loved how you threw in that comparison to enshittification, I just read the book.
Leon everything is converging! I'm a climate researcher my associate and friend also is a climate researcher however his expertise is whole system integration and analysis. He too has a substack...Regeneration. He could not help but see a significant pattern emerging with our economy and with Trumps assistance helping to rush us over a deep cliff
Here is the link to Ali's substack article:
https://ehadnameh.substack.com/p/the-convergence-how-silver-market
It might also be when you read widely enough you can see commonalities in everything!
I'll try to have a read of the link you sent, it's a long one. I've been following a great substack called System Failure by Nathan Knopp and it's basically covered history from why the Roman Empire collapsed, the rise of Christianity as the new Empire, the collapse of their authority after the black death plague, the rise of Science as the new Empire after enlightenment, etc etc. Nathan only writes a short 5 - 10 minute read on each essay so easy to digest and get through, it's very interesting and makes what's happening now more understandable, only this time civilisation is globe-spanning.
Its also possible to skim through the most important points of this. He actually just released a followup article as well. I think that the followup lends itself easier to skim most of the salient points and he too refferences histories similarities as empires rise and fall.
https://ehadnameh.substack.com/p/65e-the-seven-stages-of-systemic?utm_source=post-email-title&publication_id=2373716&post_id=182833110&utm_campaign=email-post-title&isFreemail=true&r=2ddkm6&triedRedirect=true&utm_medium=email
Thank you, I also like this connection. From my perspective, enshittification is strongly linked to a declining surface-to-volume ratio. Consumers are at the surface, and they receive progressively less attention and poorer services. The same applies to large animals: the environment becomes something external, and they are no longer able to regulate it.
This is an incredible set of active models allowing reflection on the larger systems, very thought provoking. Looking forward to reading more as further development of the model(s) you are connecting this readership to appears here in print.
How do we know that the shared metabolic rate isn't simply the convergent property of organisms jamming themselves as close as they can get to the ceiling for dissipating waste heat before their proteins lyse without going over?
Proteins start to break down at fairly high temperatures (around 40 °C), but all the organisms shown in the first figure live well below that range (most are measured at 25 deg C) and yet display similar metabolic rates. That makes it unlikely that the shared metabolic rate simply reflects a hard thermal failure limit.
Instead, it looks like the largest animals had to become endothermic to compensate for the size-related drop in mass-specific metabolic rate by generating extra heat and to catch up with the rest of the biosphere. Without doing that, they would fall outside the optimal metabolic range and would likely be at a competitive disadvantage. This is, by the way, a strong argument in favor of dinosaurs being endothermic, something that used to be considered as a heresy but gradually accepted.
See also this comment above https://bioticregulation.substack.com/p/the-small-and-the-big-lifes-fundamental/comment/192084926
Thank you Anastassia, another great read. I particularly like the sentence near the end of the article 'Trees are not a resource'. I totally agree.
Anastassia, many thanks for yet another thought-provoking article; it kept my mind busy since its publication. A couple of thoughts.
First, the consumed shared productivity over the characteristic length of a body is some kind of x^-k pattern (k being some real number). This makes sense to me for a constant input power through the sun (perhaps of the order of 200-250W/m^2 at the ocean surface) and conversion of that energy to biomass by the smallest entities, e.g. phytoplankton, at about 0.5-1W/m^2. Only a fraction of that generated stock can sustainably be ingested by larger bodies at a certain conversion rate or efficiency. That said, having a tally of all organism in size bins will allow to determine k, something > 0. However, Helen Chersky posits in her book Blue Machine that the total biomass is constant across different sizes. With a constant relationship between body size and energy consumption, k would then be rather 0; every size category would need to consume the entirety of the next smaller category, which does not sound plausible (unless it happens once). What are your thoughts on that?
Second, one of your graph shows that humanity consumes about 10% of the energy share where a more appropriate sustainable share would be sub 1%. It follows that the human population is about one to two orders of magnitude to large, or will be once the fossil energy bonanza is finished.
Thank you, Peter, for these thoughtful comments. The claim of constant total biomass is usually based on estimates of a scaling exponent that is statistically indistinguishable from zero (or -1), obtained from a linear regression of log-transformed biomass (or population density) against log-transformed body mass.
For consumed productivity, a decline of about two orders of magnitude over at least four orders of magnitude in linear size—and twelve orders of magnitude in body mass—would imply a biomass–body-mass scaling exponent of −0.17 (or, for population density against body mass, -1.17) across the full range. Because this range is rarely considered in its entirety, such a trend can easily be obscured by limited or noisy data. Thus instead of extrapolating from allometric analysis, a more robust method is a direct calculation of energy consumption for specific classes of consumers, as I did in the post and we did in our publications.
Furthermore, as we discussed in this publication https://bioticregulation.ru/common/pdf/ec04.pdf , there is evidence that biomass distributions are steeper in more stable ecosystems than in less stable ones, a factor that is almost universally ignored.
Regarding your second thought, you are right.
Anastassia, thanks for your detailed response; I'll study the paper and other literature, considering the likely relationship of body mass is probably proportional to body size to the power of three, or thereabout.
As for the second part, the consequence of a sustainable human population of about say two orders of magnitude less than would also mean that even a transition along a logistics curve towards stabilised population is unlikely (SW in LtG, 1972); rather a contraction to that stabilised world is to be expected.
Thank you for this excellent and thought-provoking writing about energy consumption. I need a little more explanation about your comment near the end of your piece: "Trees are not a resource." Since you stated what they are not, I really hoped your next statement would succinctly say what they are. This will help readers like me more easily share some concrete examples of what trees are. For example, are they biotic regulators? It seems, from your research, that trees are central to the biotic pump concept, particularly when they are part of an intact forest ecosystem. Does your statement that they are 'not a resource' argue for thinking of trees as much more than something to be exploited? Or, am I misunderstanding a major point in your writing?
Thank you for this feedback—much appreciated. Yes, you are right: trees, when they are part of a natural ecosystem, are major biotic regulators and an integral component of the biotic pump. They are, quite literally, the basis of life on land.
At the same time, I feel that these words are still too weak to fully express what I mean. That is why I deliberately left the phrase “trees are not a resource” standing on its own. My hope was that, rather than offering a ready-made conclusion, it would prompt readers to pause and think for themselves—something like opening a gestalt.
Sometimes, when a picture is complete, it appears and disappears just as quickly. But when something feels unresolved, it can linger and quietly irritate the mind for much longer. With my writing, I want to encourage people to think about the natural world in new ways.
Maybe nothing is a Resource. In 19th century classical industrial capitalist economics in which our society is still largely based, this term means "of exploitable nature" (rather than the previous meaning of an "inner strength"; Hausmann, Perreaux 2020), and to "allocate scarce means to *satisfy unlimited wants*" . This sounds like a delusional worldview resulting, perhaps, from a misplaced perception over our role on the planet. The world does not belong to us ! Life does not belong to us ! We can use what we need within balance with every other planetary inhabitant, and we are now in a better position to understand our limits to growth in both an economical and ecological sense. Unlike other species, we were able to overextend our population but now more able to keep it in check through awareness and nature-wise decisions.
Thank you for this. I'm reminded of a 2007 study in PNAS that found Human Appropriation of Net Primary Productivity is 83% in agricultural areas, and about 28% globally (that's a terrestrial fraction, I assume). That deeply brought home to scale of the ecological crisis to me.
There are also areas from which we do not consume primary productivity, but which we have anyway destroyed. This is a separate count. E.g. if you cut down trees for development, then there are no more trees which you could cut, so on this area you won't consume anything. Yet it will be destroyed and not contributing to climate stability. So there are unfortunately multiple metrics.
Absolutely. Thank you.
If you haven’t already, you should read some of Ernst Götsch’s writing on this subject. In his view, animals appear in contexts where the local net primary production has exceeded local metabolic capacity and the size of the animals is proportional to that excess. This allows for living systems to continually increase their production over evolutionary time and create larger stocks of “natural capital”. Of course these large movers of energy need predators to manage them and ensure they are working, in service as it were, to the greater system. Far from destroying vegetation, well managed large herbivores can dramatically increase NPP through a variety of mechanisms.
I would also add that large animals play a critical role in the creating more land. When large numbers of large animals are living, continental erosion rates increase (this happened with dinosaurs and Pleistocene mammals), but not above soil creation rates. Because of isostacy this means that the continent actually expands and more fertile land is created where plants can grow (erosion creates fertile deltas and fluvial depositions etc).
Interesting. Makes me remember a visit to farmers in Cameroon Highlands in Malaysia. I was horrified by the level of erosion by their farming of steep slopes. But one farmer explained to me that they got a much bigger and fertile valley bottom as a result. Also in Japan I remember similar expressions by farmers: "we cultivated away a hill"....
dear Anastassia, thanks for your contributions. I hope you keep going! In a paper from 2014 Haberl,Helmut, Karl-Heinz Erb och Fridolin Krausmann Human Appropriation of Net Primary Production: Patterns, Trends, and Planetary Boundaries, Annual Review of Environment and Resources Volume 39, 2014. estimated what they define as Human Appropriation of Net Primary Production (HANNP) to be 25 percent by 2005. Are you familiar with their calculation and if so, can you explain the difference between their estimate and the estimate by Viktor Gorshkov, apart from the fact that they are about the situation at a different times?
Thank you very much, Gunnar. I also hope to keep going, even if it has been a major shift in activities for me which I am still learning how to handle.
To compare Table 1 for HANPP from Haberl et al. 2014 with Gorshkov's estimates [https://bioticregulation.ru/common/pdf/ufn80-en.pdf, Section 11D ] it is useful to keep in mind that 1 PgC/year = 1 GtC/year = 10^12 Watt = 1 TW.
Gorshkov divided HANPP in two parts, one is power consumption due to human food, cattle fodder and timber, around 5 TW or 5 PgC/year, or 8% of terrestrial NPP (which he estimated at 60 PgC/year). This "steady-state" consumption is what is shown in the graph, to compare it with steady-state consumption in natural terrestrial ecosystems. This shows that there is an anthropogenic disturbance to stable architecture of energy flows even if the currrent food and sylviculture systems were "sustainable" in the short term.
The other part of HANPP is the decline in the biomass of the biosphere due to human disturbance (deforestation), which he estimated at 16 TW, or 27% of terrestrial NPP (giving a total of 35% for 60 GtC/year terrestrial NPP).
Vitousek et al. on which Haberl et al. are based considered not actual NPP but some potential NPP (about 75 GtC/year).
One more question: It seems as if your diagram on metabolic rate is contrary to the "mouse to elephant" curve as developed by Kleiber and others. Or do I misunderstand you or them?
https://onlinelibrary.wiley.com/doi/10.1002/j.2040-4603.2023.tb00258.x
The "mouse to elephant" curve is confined to the body size interval between the mouse and the elephant, and you can actually see it in the graph ("endotherms", also includes birds with a roughly similar scaling).
What matters is that this curve does not extend to all life, but breaks as one goes from one clade to another, and breaks in such a manner that every taxon approximately remains within the same interval of mass-specific rates.
See also this response below https://bioticregulation.substack.com/p/the-small-and-the-big-lifes-fundamental/comment/192369950
Very interesting analyses and observations. However, as far as I can analyse, Max power still applies , following the original definition by Lotka: “evolution in a system aims to increase the total (bio) mass in a system, as well as to increase the flow of mass through that system, as well as the energy flux by increasing the system, as long as there is an unused share of mass and energy in that system. ”
And growth of power ends ( but not the principle) when equilibrium is achieved, which is a result of trial and error, growth and decrease of species follows availability and dependence of each other .
Humans finding fossil stocks and ways to use these disturbed the situation and fueled max power again. Until running out of fuel and collapse.
(By the way: and humans as well shaped a imaginary system of value by money , precisely to maximize power over energy and resources. )
This is someting that ancient Taoist master Zhuang Zhou (aka. Zhuangzi, Chuang Tzu) discussed 2300 years ago in a famous story about "Man who despised machine". When I first red the story it completely changed my life. In that story Zhuang Zhou argues that in using machines as prosthetics for resource overconsumption humans selfdestruct. What impressed me is the way that Zhuang Zhou relates machine use with morality. Wise man in the story explicitly said that machine use (even simple machines) causes overexpoitation of resources. That is a kind of cheating of Nature. He knows that work is easier done with machines but he doesn't want to use them because it is immoral. Taoists know that resource overexploitation is disturbing the energy flows of nature and that means big trouble for the menkind. Overconsumption brings arrogance and arrogance brings selfdestruction. Unfortunately, when you have competitor that overexploits nature (USA) and that is arrogant and aggressive, you don't have a choice but to follow the same path in order to survive. Your only chance is to build even more powerfull machines.