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Multilevel Selection
Theoretical Foundations,
Historical Examples,
and Empirical Evidence
Multilevel Selection
Theoretical Foundations, Historical
Examples, and Empirical Evidence
Steven C. Hertler Aurelio José Figueredo
College of Saint Elizabeth University of Arizona
Morristown, NJ, USA Tucson, AZ, USA
Mateo Peñaherrera-Aguirre
University of Arizona
Tucson, AZ, USA
© The Editor(s) (if applicable) and The Author(s), under exclusive licence to Springer Nature
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Preface: A Pan-Selectionist Model
of Evolution and Development
Introduction
Open an atlas, turn the page to a map of a city, and find, in some far
corner, a smaller map of the nation containing that city. This provides
necessary perspective. While viewing a map of London, for instance, the
page will be dominated by the Thames meandering north and south as it
pushes east; also prominent will be labyrinthine streets, designated
municipal buildings, parks, landmarks, regions, and districts. Again,
however, somewhere high or low, in this corner or that, there may well be
a miniature map of the United Kingdom, on which London appears as a
dot in the Southeast of England. This introductory chapter is much like
that small map. It provides necessary perspective for the 12 content chap-
ters that follow. While this book is about multilevel selection, this intro-
ductory chapter is about evolution more generally. However, multilevel
selection progresses from possible to exceedingly plausible when contex-
tualized within the overarching pan-selectionist model of evolution pre-
sented in this preface. This pan-selectionist view of evolution permeates
boundaries between evolutionary and developmental processes within
individuals, while yoking cultural to biological evolutionary change.
Evolution is taking place, not only across generations of organisms but
within organisms and among groups of organisms, hence the need to
v
vi Preface: A Pan-Selectionist Model of Evolution and Development
preface selection with pan, which is of course a Greek prefix meaning all.
By this preface’s end, one will see how this pan-selectionist theory of evo-
lution and development is, in effect, an integrative meta-theoretical
framework in which groups are classed as one among several hierarchi-
cally nested levels of biological organization. For all this, pan-selection-
ism has precedents in the past, to which we will first turn.
Historical Precedence
Several decades ago, B. F. Skinner and E. O. Wilson noted that the object
of study in the fields which they founded, Radical Behaviorism and
Sociobiology, respectively, consists exclusively of organismal characteris-
tics, developed through the lifespan in response to genetic processes gov-
erning development coupled with one’s experiential history with an
environment (Naour, 2009). In other words, the environment acts on the
organism through the medium of genes and genetic change. Thus, ulti-
mately, genetic processes influence all characteristics selected through
development. Less commonly discussed, but also recognized by Skinner
and Wilson, is that phenotypes selected during development can have
their genetic regulation modified over evolutionary time, ultimately pro-
ducing changes in phenotypic plasticity. Thus, processes of selection by
consequences are pivotal to understanding the evolution and development
of behavior. Indeed, the common currency of selection unites an inte-
grated conception of behavioral evolution and development (e.g., Naour,
2009). Based on that conceptualization, the objective of the present pref-
atory essay is to unify our theories of evolution and development under
the overarching framework of a pan-selectionist logic.
It is important here to understand that Darwin (1859) used the word
“selection” as an anthropomorphic metaphor personifying “Nature” as an
intentional agent. In reality, all that happens is the differential survival
and reproduction of alternative variants. Darwin was quite clear that he
used this expression as just a figure of speech:
to avoid personifying the word Nature; but I mean by Nature, only the
aggregate action and product of many natural laws, and by laws the
sequence of events as ascertained by us. (p. 81)
The 10,000 or so synapses per cortical neuron are not established immedi-
ately. On the contrary, they proliferate in successive waves from birth to
puberty in man… One has the impression that the system becomes more
and more ordered as it receives ‘instructions’ from the environment. If the
theory proposed here is correct, spontaneous or evoked activity is effective
only if neurons and their connections already exist before interaction with
the outside world takes place. Epigenetic selection acts on preformed syn-
aptic substrates. To learn is to stabilize preestablished synaptic combina-
tions, and to eliminate the surplus. (p. 248)
One such simple social norm might be stated as follows: “Stop on red,
go on green, exert caution on yellow,” in reference to traffic signals.
Within that rule is embedded an implicit verbal statement in the form of
a syllogism: “if A–and–if B–then C,” pointing to contingent relations
among Antecedent conditions, target Behaviors, and their probabilistic
Consequences. For example, if the light is red, and if you do not stop,
then the police will issue you a traffic ticket, associated with a monetary
fine or penalty. The monetary fine or penalty, however, is merely the
proximal or social consequence of nonadherence to the social norm.
Behind it is a more distal or natural consequence of nonadherence, one
of possibly being involved in a collision with another moving vehicle—
one that presumably is adhering to the norm.
Preface: A Pan-Selectionist Model of Evolution and Development xiii
The genetic assimilation of social norms could thereby give rise to the
phenomenon of biocultural group selection, wherein different biocul-
tural groups might be discriminable both culturally and genetically based
on their divergently evolved normative structures and might thus be
selected among as higher-order vehicles for their inner replicators (see
Sarraf, Woodley of Menie, & Feltham, 2018). Thus, the epigenetic rules
of development controlling gene-culture coevolutionary processes reflect
systematic and lawful patterns of gene-environment interaction, ulti-
mately controlled by the molecular biology of environmentally contin-
gent gene expression. It becomes increasingly apparent that these various
processes of selection by consequences at the phenotypic level are more
than merely analogous, but turn out to be homologous, in the full evolu-
tionary sense of the word.
(that they are thus able to “recognize”), and to which they produce epige-
netically variant specific antibodies, are selectively proliferated. This
occurs via two different causal pathways: (1) the B-lymphocyte cells
encountering the antigens are themselves triggered to proliferate by
increased rates of mitosis; and (2) these same cells send paracrine chemi-
cal signals to the pluripotent hematopoietic progenitor cells in bone mar-
row, prompting them to produce more of the same specific variety of
epigenetically variant lymphocyte. There appear to be no mechanisms
selecting against other types of variants, which may continue to persist in
their previous numbers, albeit in reduced proportions with respect to the
former as they are not proliferated.
In the nervous system, on the other hand, the selection among pro-
cesses can be best characterized as selection against certain types of vari-
ants, in that any epigenetically modified neurons that do not fire at a rate
above their base rates are selectively eliminated by the selective with-
drawal of a transcription factor called nerve growth factor (NGF) that is
needed for their continued survival. A transcription factor is a protein
that binds to the target DNA molecule and either upregulates or down-
regulates the synthesis of its specific gene product. Withdrawal of NGF
thereby activates the mitochondrial pathway of apoptosis (among others)
and results in the selective and programmed cell death of the underper-
forming neurons. In this case of selection against, there appears to be no
selection for other epigenetically variant neurons, as they may continue to
persist in their previous numbers, albeit in increased proportions with
respect to the ones eliminated, but they are not in any way proliferated by
this process.
These considerations all converge upon the logical implication that
selection by consequences at the phenotypic level is ultimately regulated
biologically by means of genetic and epigenetic mechanisms of develop-
ment and thus subject to adaptive evolution. For example, biological pre-
paredness is the evolved genetic programming of developmental switching
mechanisms that bias development disproportionally toward alternative
developmental outcomes that, in the case of reinforcers, influence which
stimuli in the environment do and do not have reinforcing or punishing
properties relative to individual behavior and to the relative degrees they
each possess such values.
Preface: A Pan-Selectionist Model of Evolution and Development xvii
SUN CHIA-NAI
This official, chiefly known to fame among his countrymen as one
of the tutors of His Majesty Kuang-Hsü, was a sturdy Conservative of
the orthodox type, but an honest and kindly man. His character and
opinions may be gauged from a well-known saying of his: “One
Chinese character is better than ten thousand words of the
barbarians. By knowing Chinese a man may rise to become a Grand
Secretary; by knowing the tongues of the barbarians, he can at best
aspire to become the mouth-piece of other men.”
Ceiling and Pillars of the Tai Ho Tien.
In his later years he felt and expressed great grief at the condition
of his country, and particularly in regard to the strained relations
between the Empress Dowager and the Emperor. He traced the first
causes of these misfortunes to the war with Japan, and never
ceased to blame his colleague, the Imperial Tutor Weng T’ung-ho,
for persuading the Emperor to sign the Decree whereby that war was
declared, which he described as the act of a madman. Weng,
however, was by no means alone in holding the opinion that China
could easily dispose of the Japanese forces by land and by sea. It
was well-known at Court, and the Emperor must have learned it from
more than one quarter, that several foreigners holding high positions
under the Chinese Government, including the Inspector-General of
Customs (Sir Robert Hart), concurred in the view that China had
practically no alternative but to declare war in view of Japan’s high-
handed proceedings and insulting attitude. Prestige apart, it was
probable that the Emperor was by no means averse to taking this
step on his own authority, even though he knew that the Empress
Dowager was opposed to the idea of war, because of its inevitable
interference with the preparations for her sixtieth birthday; at that
moment, Tzŭ Hsi was living in quasi-retirement at the Summer
Palace. After war had been declared and China’s reverses began,
she complained to the Emperor and to others, that the fatal step had
been taken without her knowledge and consent, but this was only
“making face,” for it is certain that she had been kept fully informed
of all that was done and that, had she so desired, she could easily
have prevented the issue of the Decree, and the despatch of the
Chinese troops to Asan. Sun Chia-nai’s reputation for sagacity was
increased after the event, and upon the subsequent disgrace and
dismissal of Weng T’ung-ho he stood high in Her Majesty’s favour.
Nevertheless his loyalty to the unfortunate Emperor remained
unshaken.
In 1898, his tendencies were theoretically on the side of reform,
but he thoroughly disapproved of the methods and self-seeking
personality of K’ang Yu-wei, advising the Emperor that, while
possibly fit for an Under-Secretaryship, he was quite unfitted for any
high post of responsibility. When matters first approached a crisis, it
was by his advice that the Emperor directed K’ang to proceed to
Shanghai for the organisation of the Press Bureau scheme. Sun,
peace-loving and prudent, hoped thereby to find an outlet for K’ang
Yu-wei’s patriotic activities while leaving the Manchu dovecots
unfluttered. Later, after the coup d’état, being above all things
orthodox and a stickler for harmonious observance of precedents, he
deplored the harsh treatment and humiliation inflicted upon the
Emperor. It is reported of him that on one occasion at audience he
broke down completely, and with tears implored the Empress
Dowager not to allow her mind to be poisoned against His Majesty,
but without effect.
Upon the nomination of the Heir Apparent, in 1900, which he, like
many others, regarded as the Emperor’s death sentence, he sent in
a strongly worded Memorial against this step, and subsequently
denounced it at a meeting of the Grand Council. Thereafter, his
protests proving ineffective, he resigned all his offices, but remained
at the capital in retirement, watching events. At the commencement
of the Boxer crisis, unable to contain his feelings, he sent in a
Memorial through the Censorate denouncing the rabid reactionary
Hsü T’ung, whom he described as “the friend of traitors, who would
bring the State to ruin if further confidence were placed in him.”
Throughout his career he displayed the courage of his convictions,
which, judged by the common standard of Chinese officialdom, were
conspicuously honest. He was a man of that Spartan type of private
life which one finds not infrequently associated with the higher
branches of Chinese scholarship and Confucian philosophy; it was
his boast that he never employed a secretary, but wrote out all his
correspondence and Memorials with his own hand.
A pleasing illustration of his character is the following: He was
seated one day in his shabby old cart, and driving down the main
street to his home, when his driver collided with the vehicle of a well-
known Censor, named Chao. The police came up to make enquiries
and administer street-justice, but learning that one cart belonged to
the Grand Secretary Sun, they told his driver to proceed. The
Censor, justly indignant at such servility, wrote a note to Sun in which
he said: “The Grand Secretary enjoys, no doubt, great prestige, but
even he cannot lightly disregard the power of the Censorate.” Sun,
on receiving this note, proceeded at once on foot in full official dress
to the Censor’s house, and upon being informed that he was not at
home, prostrated himself before the servant, saying: “The nation is
indeed to be congratulated upon possessing a virtuous Censor.”
Chao, not to be outdone in generosity, proceeded in his turn to the
residence of the Grand Secretary, intending to return the
compliment, but Sun declined to allow him to apologise in any way.
TUAN FANG
In 1898, Tuan Fang was a Secretary of the Board of Works; his
rapid promotion after that date was chiefly due to the patronage of
his friend Jung Lu. For a Manchu, he is remarkably progressive and
liberal in his views.
In 1900, he was Acting-Governor of Shensi. As the Boxer
movement spread and increased in violence, and as the fears of
Jung Lu led him to take an increasingly decided line of action against
them, Tuan Fang, acting upon his advice, followed suit. In spite of
the fact that at the time of the coup d’état he had adroitly saved
himself from clear identification with the reformers and had penned a
classical composition in praise of filial piety, which was commonly
regarded as a veiled reproof to the Emperor for not yielding implicit
obedience to the Old Buddha, he had never enjoyed any special
marks of favour at the latter’s hands, nor been received into that
confidential friendliness with which she frequently honoured her
favourites.
In his private life, as in his administration, Tuan Fang has always
recognised the changing conditions of his country and endeavoured
to adapt himself to the needs of the time; he was one of the first
among the Manchus to send his sons abroad for their education. His
sympathies were at first unmistakably with K’ang Yu-wei and his
fellow reformers, but he withdrew from them because of the anti-
dynastic nature of their movement, of which he naturally
disapproved.
As Acting-Governor of Shensi, in July, 1900, he clearly realised
the serious nature of the situation and the dangers that must arise
from the success of the Boxer movement, and he therefore issued
two Proclamations to the province, in which he earnestly warned the
people to abstain from acts of violence. These documents were
undoubtedly the means of saving the lives of many missionaries and
other foreigners isolated in the interior. In the first a curious passage
occurs, wherein, after denouncing the Boxers, he said:
After prophesying for them the same fate which overtook the
Mahomedan rebels and those of the Taiping insurrection, he
delivered himself of advice to the people which, while calculated to
prevent the slaughter of foreigners, would preserve his reputation for
patriotism. It is well, now that Tuan Fang has fallen upon evil days, to
remember the good work he did in a very difficult position. His
Proclamation ran as follows:—
“If the rain has not fallen upon your barren fields,” he said,
“if the demon of drought threatens to harass you, be sure that
it is because you have gone astray, led by false rumours, and
have committed deeds of violence. Repent now and return to
your peaceful ways, and the rains will assuredly fall. Behold
the ruin which has come upon the provinces of Chihli and
Shantung; it is to save you from their fate that I now warn you.
Are we not all alike subjects of the great Manchu Dynasty,
and shall we not acquit ourselves like men in the service of
the State? If there were any chance of this province being
invaded by the enemy, you would naturally sacrifice your lives
and property to repel him, as a matter of simple patriotism.
But if, in a sudden access of madness, you set forth to
butcher a few helpless foreigners, you will in no wise benefit
the Empire, but will merely be raising fresh difficulties for the
Throne. For the time being, your own consciences will accuse
you of ignoble deeds, and later you will surely pay the penalty
with your lives and the ruin of your families. Surely, you men
of Shensi, enlightened and high-principled, will not fall so low
as this? There are, I know, among you some evil men who,
professing patriotic enmity to foreigners and Christians, wax
fat on foreign plunder. But the few missionary Chapels in this
province offer but meagre booty, and it is safe to predict that
those who begin by sacking them will certainly proceed next
to loot the houses of your wealthier citizens. From the burning
of foreigners’ homes, the conflagration will spread to your
own, and many innocent persons will share the fate of the
slaughtered Christians. The plunderers will escape with their
booty, and the foolish onlookers will pay the penalty of these
crimes. Is it not a well-known fact that every anti-Christian
outbreak invariably brings misery to the stupid innocent
people of the district concerned? Is not this a lamentable
thing? As for me, I care neither for praise nor blame; my only
object in preaching peace in Shensi is to save you, my
people, from dire ruin and destruction.”