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Dear Mats, dear colleagues,Thank you very much for continuing the discussion. Below is my take on Mats' comments. In general, I would like to emphasize the need for quantitative rather than qualitative arguments to differentiate between biotic regulation versus adaptation (see my concluding commentary below). Finding such arguments constitutes the essence of the biotic regulation scientific enterprise.Some natural forests are very old. In Australia I was told that the rainforest in the northeast was 100 million years old. Even if this perhaps was an exaggeration, it is certainly a very old forest. It must have lived through periods with carbon dioxide concentrations much higher than today. If the biota has been controlling the carbon dioxide concentration during this time, how come that the concentration has changed so much? What has made the biota change the optimal concentration? I think it is much more likely that other factors have made the climate change, and that the biota has adapted. It is controlling as well as it can, but not better.
"Changed so much" -- compared to what? Compared to the biotic sensitivity. One can argue that the environmental changes that did occur under the influence of abiotic factors are of a smaller amplitude than they would have been in the absence of the biotic regulation. I do realize that the biota has a strong regulatory effect. The biota is certainly controlling as well as it can. This control is apparently efficient enough to keep the environment liveable despite the influence of the many destabilising abiotic factors. And, from the fact that the environment nevertheless undergoes certain changes it does not follow that the biota is adapting. The biota is surviving through the less optimal conditions and is, in the meantime, continuing to push the environment back to the optimum. What I wonder is the level of the optimum. According to the results of Royer et al. (https://doi.org/10.1038/ngeo1186) on the variation of CO2 over cenozoicum, the values were in the range 800 and 2000 ppm (with large uncerainties) between 52 and 32 Mya, and then declined (with some ups and downs) and have stayed mainly between 300 and 180 ppm during the last 20 Mya. The optimum concentration should be fixed within the biotic sensitivity (about 1% or 3ppm according to one of your papers). So do you propose that the biota for some unknown reason has changed the optimum from somewhere in the order of 1000 ppm to maybe 250ppm, or has it been pushing incessantly for 40 My? Maybe it is pushing now towards a high optimum?
An ecological community in one part of the world must be exposed incessantly to a stream of alien organisms, in particular spores, bacteria, viruses and so on, but also occasionally larger organisms (with their associated microbiomes). Many will probably be fended off, but some will be introduced, may be after an initial epidemic period. This will change the composition and the genetic program of the community. It cannot be so rigorously specified as point 2 above asserts.
Again, it is difficult to discuss such issues outside a quantitative context. "Some will be introduced" -- this can be a very rare event that is comparable in frequency to the origin of a new species in the course of evolution. (This occurs about once in a few million years within each lineage). A few million years also for bacteria?Point 2 in your quotes refers to the distribution of energy fluxes over organisms of different sizes. It is universal in stable ecosystems on land and in the sea, with the consumption by the largest organisms strictly limited (albeit by different mechanisms) (see Nefiodov 2020 and Fig. 4 here ). That was an important clarification! My quote was …each community is characterised by 1) a strictly- specified species composition and 2) a strictly-specified distribution of fluxes of matter and energy going through different species in the community. I should replace "species" by "size-ranges of species" or something like that? Should I do so also in the first part? Then I have no objections. I think that the question about how large (strictly is not quantitative statement!) a variation of species there is, in particular in the microbiome, is a question for future research. The "everything is everywhere" statement seems to be questioned more and more. See https://doi.org/10.1073/pnas.1212424110.
During the last 2.75 Myr periods of glaciations have been interrupted by brief interglacials, which I think can be basically understood from the Milankovitch cycles, even if many feedback mechanisms are but poorly understood. The importance of biotic regulation as a negative feedback has not been properly recognized. But I do not believe in the proposition that the glaciations are primarily due to disruption of the biota by big animals. After all, the distribution of sunlight over the planet can not be controlled by the biota. Over a cycle of about 20 000 years northern latitudes change from a period with short cool summers, to one with long warm summers. Vegetation zones are moving south and north in sync with this cycle.
Glaciation is associated with a drop in the global mean surface temperature. If our planet were covered by ocean, with no hemispheric land/ocean asymmetry, the changes of sunlight distribution between the hemispheres could not have produced glaciations -- warming in one hemisphere would have been offset by cooling in the other. So the fact (if we accept it as a fact) that the periodic changes in sunlight distribution among the hemispheres triggers glaciations, implies that there is a positive feedback that amplifies the temperature changes in one hemisphere but not in the other. With most land on the nothern hemisphere and mainly ocean on the southern, it is easy to see that a lot of physical feedback mechanisms would set in. The changing vegetation cover is of course very important, and probably also big herbivores. It is again a question where a quantitative context would be useful!
Given that the terrestrial vegetation cover exerts a profound influence on both albedo and the greenhouse effect on the continental scale, and that the big herbivores have the potential to massively destroy the vegetation cover, I consider it plausible that without the big herbivores the destabilising positive feedbacks would have been weaker or even absent at all. The forests (and other ecosystems like tundra) could have then possibly survived the short cool summers as part of the climate variability that they are programmed to be able to endure. The amplitude of global cooling could have been smaller and glaciations less pronounced.
Note also that the fact that once the glaciations do occur and retreat, the vegetation zones are moving south and north, shows that the corresponding ecosystems do not adapt. Indeed, the timescale of glaciations (ten to hundred thousand years) is much less than the characteristic timescale of biological evolution (several million years). This is true. But on one geographical spot the ecosystem changes from one (e.g. forest) to another (e.g. tundra) in a continuous(?) fashion. This sort of adjustment only involves a presumably concerted change of species in the ecosystem, so its is not an adaptation genetically. I do not know if tundra is a stable ecosystem, but I can imaging a similar change over from one type of stable system to another. Would such a change always involve a discontinuous break, destruction of the first type and succesive establishment of the second?
Forest areas, steppes, deserts have waxed and waned. When a forest system has returned to a northerly area after a glaciation, it will establish itself on a soil that is very different from that before the glaciation. Will the ecological communities before and after be exactly the same?
Yes they will. After any fire, cutting or even just a big old tree falling down, the soil conditions change dramatically. Nevertheless, the ecological community is equipped with all the needed species that will initiate succession and on the time scale of a few hundred years restore the unperturbed climax community. E.g., colonization of a bare rock can be started by lichens that form an organic layer where other species can later establish. So a post-glaciation environment does not present any particular variability compared to what the ecological community normally experiences during succession. i am again thinking on the microbiome. In this case the ubiquity hypothesis is probably applicable, so it is a question about how the numbers of dominant species varies in the microbiome, and how long it takes for the composition to return to the old state. We (or those of us that are still alive) will learn more about that in the future.
A natural forest comprises sets of communities that all exert regulation of the environment. In different parts of the earth there are very different forest systems, containing entirely different species within their communities. Each of these forests must evidently be set to regulate global variables to the same values. How can this come about, without some sort of adaptation?Given that the global conditions (e.g. the CO2 concentration) are about the same to all, it is clear that all ecosystems should work to maintain one and the same value for all global parameters. I see nothing surprising in this.Notably, life originated in the ocean. Without any life on land, the oceanic biota had been able to keep the environment stable. When the terrestrial biota began to emerge, it was local and had to conform to the global and regional conditions kept by the oceanic biota. So some of the global conditions (e.g. oxygen concentrations) remain beyond the control of the terrestrial biota and are regulated by the ocean (see discussion in Section 4.2 here).[biotic regulation] must be made compatible with the striking adaptation of living beings to their environment, including their fellow beings
There is adaptation as a state (sometimes referred to as adaption) and adaptation as a process. The state of adaptation refers to the observation that there is a strict correlation between properties of different organisms that are essential for their survival. This correlation is not just compatible with biotic regulation but constitutes the essence of it. The various species are correlated to each other as are the parts of a complex machine. The properties of the machine as a whole are correlated with the properties of the environment where it works and which it controls. It cannot be otherwise. The biotic regulation mechanism may evolve, but this evolution will preserve the correlation between its parts. I am still not sure about what you are aiming at. To attain a state of adaptation is a process that I would call adaptation. Example: For flowers and pollinaters to become so exceedingly well adapted to each other, it seems to me that they have gone through a process that could be called adaptation. But I am rather ignorant also about this. I shall try to understand a little more about ecology and information, using some of the references you have provided.
Darwinian evolution as a concept is agnostic concerning the environment. It automatically identifies any evolutionary change with an adaptation (or just sees it as a stochastic process) but it does not present any arguments about how the presumed genetic adaptation is quantitatively related to environmental change. E.g. what is the probability that for a given change in an environmental parameter there will be a genetic variant found in a given population that will be able to sustainably maintain a higher viability under new conditions.
One of the key quantitative arguments against environmental adaptation is the fact that speciation proceeds at an approximately the same rate (one species in several million years) in organisms generating genetic diversity at vastly different rates (differing by many orders of magnitude) (see here ). If evolutionary changes were driven by this generation of genetic diversity, we could expect that the smaller numerous organisms (that generate a lot of diversity) would be more rapidly evolving reacting to even short-term changes in environmental conditions. This is not the case. Yes, this is a strong argument, and as I have said earlier I was very impressed by it. It is even more striking considering that the small organisms generally have short generation lenghts. Do you think that the speciation rate is reflecting the time it takes for the the whole "machine" to readjust – introduction of a new or changed species must involve a lot of readjustment. – But I am still worried about the inflexibility (it seems infexible to me) of the biotic machinery as you describe it. Life is extremely flexible. Trees in a forest grew in different habitats, depending among other things on the topology of the landscape, with different macro companions, and certainly also different microbiomes. The local regulation should work well over all different habitats, and then there must be a lot of variation at least in the numerical composition of communities that all should work well enough.
Best wishes,Anastassiaпн, 7 дек. 2020 г. в 17:50, Mats Almgren <mats.a...@gmail.com>:Dear Anastassia, dear all,I have started to slowly reread the 2000 Springer book, trying to be very critical. So far I have just begun, but even so I have a number of questions. I have extracted three assertions:
- Ecological communities exert strict (within the biotic sensitivity) control of the environment, even of global parameters such as the carbon dioxide concentration in the atmosphere.
- Ecological communities are strictly specified and invariable:…each community is characterised by 1) a strictly- specified species composition and 2) a strictly-specified distribution of fluxes of matter and energy going through different species in the community. It follows that also the relative number of individuals of different species must be fixed!
- These two reactions to environmental changes—biotic regulation and genetic adaptation—are incompatible in the sense that only one of them is actually realised. Individuals of biological species may either change the environment to the initial optimal state, or change themselves by genetically adapting to the new environment. There is no compromise between the two possibilities.
With regard to them I have the following remarks:
- A natural forest comprises sets of communities that all exert regulation of the environment. In different parts of the earth there are very different forest systems, containing entirely different species within their communities. Each of these forests must evidently be set to regulate global variables to the same values. How can this come about, without some sort of adaptation?
- Some natural forests are very old. In Australia I was told that the rainforest in the northeast was 100 million years old. Even if this perhaps was an exaggeration, it is certainly a very old forest. It must have lived through periods with carbon dioxide concentrations much higher than today. If the biota has been controlling the carbon dioxide concentration during this time, how come that the concentration has changed so much? What has made the biota change the optimal concentration? I think it is much more likely that other factors have made the climate change, and that the biota has adapted. It is controlling as well as it can, but not better.
- During the last 2.75 Myr periods of glaciations have been interrupted by brief interglacials, which I think can be basically understood from the Milankovitch cycles, even if many feedback mechanisms are but poorly understood. The importance of biotic regulation as a negative feedback has not been properly recognized. But I do not believe in the proposition that the glaciations are primarily due to disruption of the biota by big animals. After all, the distribution of sunlight over the planet can not be controlled by the biota. Over a cycle of about 20 000 years northern latitudes change from a period with short cool summers, to one with long warm summers. Vegetation zones are moving south and north in sync with this cycle. Forest areas, steppes, deserts have waxed and waned. When a forest system has returned to a northerly area after a glaciation, it will establish itself on a soil that is very different from that before the glaciation. Will the ecological communities before and after be exactly the same?
- An ecological community in one part of the world must be exposed incessantly to a stream of alien organisms, in particular spores, bacteria, viruses and so on, but also occasionally larger organisms (with their associated microbiomes). Many will probably be fended off, but some will be introduced, may be after an initial epidemic period. This will change the composition and the genetic program of the community. It cannot be so rigorously specified as point 2 above asserts.
These points suffice for the moment. In short my view is that there is a lot in favour of the biotic regulation theory. But I think that in some way it must be made compatible with the strikning adaptation of living beings to their environment, including their fellow beings. I also believe that biotic regulation theory provides a strong negative feedback to environmental changes, but not that it can control the environment so strictly. If I understand your view correctly, the planet has never been covered by stable ecosystems. Steppes, savannahs, deserts, shorelines, rivers and more are all unstable. Natural forests and the oceans try to regulate the environment; on the rest of the planet life is just going on in an unstable way, with events of overpopulation, extinctions, famines and so on. Seems like a stage for a Darwinian evolution.I appreciate any comments. I realize that most of you have more important things to do, but I think that this biotic regulation theory is very important and needs to be critically and seriously discussed.Best regardsMats
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Dear colleagues,
just a note about this hugely interesting conversation. I just
wanted to let you know that I have been following and mulling some
ideas of mine in this respect. But these ideas still have to be
refined, so for the time being, I'll be just listening.
And since I am here, I thought I could wish to all of you the
best for this end of the year, which has not been a good year, and
it is ending even worse. Anyway, we keep going and, who knows?
humankind may even learn something, one day or another.
ATB
Ugo
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-- *********************************** Prof. Ugo Bardi Dipartimento di Chimica, Università di Firenze, Italy Full member of The Club of Rome Registered Expert of the United Nations "Harmony With Nature" Programme Chief Editor of "Biophysical Economics and Sustainabilty," a Springer Journal Delegato del Rettore per la gestione della Rete Università Sostenibili (RUS) ugo....@unifi.it www.cassandralegacy.blogspot.com www.theproudholobionts.blogspot.com
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