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Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !

onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
A world of opportunity within constraint:
Pere Alberchs early EvoDevo
Arantza Etxeberria
Laura Nuo de la Rosa
1. Introduction
The work of Pere Alberch is crucial to study the early stages of Evoevo! "n #articular$
it illustrates very #ersuasively why develo#%ental syste%s have so %uch to say about
the course of evolutionary change$ since in addition to an i%#ortant e%#irical work he
elaborated a sti%ulating fra%ework of theoretical ideas on biological for%$ on the
configuration of %or#hological variation$ and on how develo#%ental #rocesses
establish #ossible evolutionary #aths #revious to the action of natural selection! Thus$
the study of develo#%ent and evolution are related through the notion of #ossible
%or#hologies! "n his view$ the %or#hology of organis%s shows internal coherence and
structure$ e%ergent fro% co%#lex non linear interactions a%ong #arts and with the
environ%ent! This constitutes a source of deter%inis% absent fro% accounts in which
novelty is considered to a##ear only fro% rando% %utations! "n his words& '"n evolution
selection %ay decide the winner of a given ga%e but develo#%ent non(rando%ly selects
the #layers' )*+,-$ #! ../0!
"n the 1-s and ,-s %any biologists argued against the #revalent gene(centred view of
biology and de%anded to study how the organization of %ulticellular organis%s$ their
Bauplan$ influences %or#hological variation and$ conse2uently$ evolutionary change! "n
Alberch3s work the link between develo#%ent and evolution is recovered in several
ways& First$ he challenged the static hierarchical theories of develo#%ent in favour of a
dyna%ic$ cyclical and interactive conce#tion that he #ursued through ex#eri%ents and
dyna%ical %odels! Second$ he stressed the relevance of studying ontogenetic
#heno%ena at the a##ro#riate %or#hogenetic level$ so that %or#hologies %ight be
regarded as #roducts of co%#lex genetic and e#igenetic interactions! Finally$ he
%aintained that so%e develo#%ental #ro#erties directionally constrain #ossible #aths of
evolution$ because they define the set of variations associated with certain for%s )e!g!$
the tetra#od li%b$ the #attern of digits or the for% of teeth0! Thus$ Alberch challenged
the theoretical fra%ework that assu%es a direct corres#ondence between genetic
structure and %or#hology$ with natural selection acting on the effects of rando%
changes!
Although Alberch3s work flourished #recisely before and after the ex#ansion of
develo#%ental genetics$ he was %ore interested in the %athe%atical techni2ues of
dyna%ical syste%s than in the new %olecular techni2ues a##earing at the ti%e! 4is
a##roach was that of a theoretical biologist who considered that ex#eri%ental and
theoretical work together could allow for an a #riori and #redictive knowledge of living
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
for% and its evolution! Thus$ Alberch5s #eculiarity with res#ect to others investigating
develo#%ent fro% a dyna%ical syste%s a##roach$ like 6oodwin )*++70 or 8auff%an
)*++90$ is the use of ex#eri%ents in addition to si%ulations!
"n his relatively short life$ Alberch wrote %any articles on %any sub:ects )he #ublished
his first #a#er when he was only *+0$ but #robably his %ost i%#ortant$ and %ost cited$
work was done during the decade of the ,-s! ;any of the advance%ents in Evoevo
these days are a conse2uence of ideas and #ro:ects in which he$ along with %any others$
was involved at that ti%e!
"n this #a#er we focus on what we consider to be the %ain early ideas of Evoevo! <irst
we see that Alberch defends the i%#ortance of doing research at the %or#hological
level$ which he considers as non reducible to lower level %olecular factors and that
a##ears as a real% of discrete the%es )section =0! Then$ we focus on the relevance of
develo#%ental %echanis%s and the role they ac2uire for studying evolution )section 90
and the way those %echanis%s are conce#tualized as develo#%ental constraints to
reflect the intrinsic abilities of develo#%ental syste%s to #roduce so%e for%s and not
others$ and the conse2uences for evolutionary o##ortunities )section 70! <inally$ in the
last section we su%%arize so%e conce#tual ele%ents of Alberch>s views on how to
integrate different a##roaches in order to ex#lain the %or#hological organization of
living beings!
2. The themes of the discrete morphospace
The #heno%enological de#arture #oint of Alberch5s research is the recognition of the
ordered character of %or#hos#ace$ i!e! the s#ace of biological for%s& ?Nature is not
chaos$ neither is it a boundless continuu% of for%s!@ )*+,=$ #! 9*/0! ;or#hological
variation is not rando% and continuous$ as the standard view in evolutionary biology
%aintain$ but co%es in ?the%es@&
)*0 Phenoty#es are discrete! That is$ #oints are not distributed unifor%ly over
#henoty#e s#ace$ but tend to cluster around %a:or 'the%es'$ corres#onding to taxa
or classes of teratologies! )=0 Ahile there %ay be considerable dis#ersion around a
%or#hological the%e$ the variability in any trait is definitely li%ited! )90 Ahen new
%or#hological the%es arise$ either in ontogeny or #hylogeny$ the transitions
between the%es are not rando%! )70 These #ro#erties are largely the result of
e#igenetic interactions during develo#%ent! )Bster C Alberch *+,=$ #! 7770
The neo(arwinian a##roach cannot fully account for the discrete character of
%or#hos#ace& if evolution strongly de#ends on the action of natural selection on gradual
variations$ any living for% %ust be theoretically #ossible$ and biological %or#hologies$
continuous! <ro% this #ers#ective$ discrete characters %ust be the result of functional
convergence& unity of ty#e in non(related taxa is seen as the outco%e of si%ilar
selective #ressures!
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
"n Alberch5s view$ internal )or develo#%ental0 and external )or selective0 ex#lanations
of evolutionary change are not contradictory$ but they deal with different #roble%s& the
origin and the fixation of evolutionary change! Dtandard evolutionary biology has been
traditionally concerned with the fate$ not the origin$ of variation! The neglect of the
origin of variation is linked to the beliefs of the isotro#y of %olecular variation and the
direct corres#ondence of genoty#e and #henoty#e! Alberch challenged both assu%#tions
and vindicated the study of the patterns of variation #recisely in order to understand the
%echanis%s res#onsible of their generation$ i!e! the variability of develo#%ental
syste%s$ as %or#hological #arallelis%s suggest that the sa%e %echanis%s are in action
)*+,90! Thus$ in his view$ evolutionary research needs to #roceed in two ste#s& the first
is to elucidate an ordered #attern of variation$ i!e! to characterize invariance$ and the
second$ to look for the %echanis%s behind those #atterns )*+,/b0! Aithout
characterizing %or#hological invariance (he argued( it would be i%#ossible to find the
%echanis%s res#onsible of its generation! That is why in Alberch5s work %or#hology
recovers the significance as a biological disci#line that it lost in the ;odern Dynthesis&
co%#arative anato%y has a funda%ental role in evolutionary theory$ since it is its :ob
?to deter%ine regularities of structural organization that enable a classification and
understanding of the ordered diversity of for%@ )Dhubin C Alberch *+,.$ #! 9110!
The study of the #atterns of variation is thus the first ste# of Alberch5s research and it is
also the sole #rotagonist of %any of his #a#ers$ where he tries to distinguish between
the order internally generated by develo#%ent and that one #roduced by natural
selection! Aith this goal$ Alberch studied three %ain %or#hological #heno%ena where
the #atterns of variation cannot be ex#lained by Natural Delection but by the structure of
develo#%ental #rocesses!
Bne case is that of morphological convergence where a si%ilar %or#hology a##ears
linked to different ada#tive re2uire%ents! "n a series of ex#eri%ents$ the functional
significance of structural %odifications in hand and foot %or#hology )such as reduction
and loss of #halangeal ele%ents$ develo#%ent of interdigital webbing and fusion of
tarsal and car#al ele%ents0 is ex#lored! The goal is to analyse how to integrate
ada#tation and ontogeny in a case where a %or#hological convergence does not i%#ly
functional convergence )*+,*0!
A second #heno%enon of %or#hological order is that revealed by the #atterns of
intraspecific variation! This is the sub:ect of Alberch5s study of #henoty#ic variation in
osteological characters of #o#ulations of s#ecies of the neotro#ical sala%ander genus
Bolitoglossa )*+,90! The %ain finding is a recurrence of the sa%e variants in widely
unrelated s#ecies$ a fact that again$ cannot be ex#lained fro% an externalist a##roach!
<inally$ a constant source of evidence for discrete variation co%es fro% his studies of
teratologies! Throughout his work$ Alberch )like transcendental %or#hologists Etienne
and "sidore 6eoffroy Daint 4ilaire0 de%onstrated a s#ecial fascination for the
develo#%ent of %alfor%ations )*+,+0! Non(functional teratologies were es#ecially
a##ealing for the study of the generative #ro#erties of develo#%ental syste%s! These
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
deviations of 5nor%al5 develo#%ent result in for%s which are often lethal or less well
ada#ted than its #redecessors! Therefore$ following the ada#tationist logic$ natural
selection should #revent their a##earance or at least contribute to their disa##earance!
Eut in s#ite of a strong selection against$ teratologies are still being generated in a
recurrent way!
All these evidences of %or#hological order$ all these #atterns of variation$ account for
the funda%ental role of develo#%ent in s#ecifying the #ossible for%s available for
selection& %or#hological convergence$ intras#ecific %or#hological diversification and
teratologies show a logic e%erging fro% the internal order of develo#%ental #rogra%s!
3. Discoverin developmental mechanisms
Ae have seen that the #ro#erties of %or#hos#ace de%and that %or#hological evolution
be studied fro% an ontogenetic #ers#ective )*+,-0! This re2uires to define
develo#%ental #heno%ena at the %or#hogenetic level& des#ite the standard #osition of
#o#ulation genetics according to which all evolutionary novelty co%es fro% rando%
%utations$ in a direct one(to(one genoty#e(to(#henoty#e relation$ the study of ontogeny
obliges to #ay attention to ?e#igenetic regulation@ and$ thus$ to a level higher than that
of genetics to ex#lain develo#%ent!
The %or#hogenetic level is characterized in two ways! Bn the one hand$ develo#%ental
for% is not a static entity defined by the s#atial #osition of its co%#onent #arts$ but the
result of a dynamical #rocess involving interactions a%ong the%$ integration and
regulation! Bn the other hand$ the %or#hogenetic level %ay be studied in #hysico(
che%ical ter%s! These two #oints converge in a conce#tualization of develo#%ental
dyna%ics as susce#tible of %athe%atical for%alization and ex#eri%ental treat%ent!
This section will #resent Alberch5s conce#tualization of the %or#hogenetic level and the
global #ro#erties of develo#%ental #rocesses! Next$ both work on the %athe%atical
for%alization of the dyna%ics of develo#%ental syste%s )and the role of %athe%atical
biology0 and of their bio#hysical #ro#erties )and the role of ex#eri%ental e%bryology0
are considered! Eoth a##roaches define a %echanistic view#oint on develo#%ent and
evolution that %ay #rovide so%e degrees of deter%inis% and #redictability!
3.1. The morphogenetic level
Bne of the %ain difficulties for Alberch5s #osition was to establish the reality of the
%or#hogenetic level and the necessity of working at it des#ite the success of the
geno%e(centred a##roach in evolutionary and develo#%ental theory! "n his o#inion$ the
latter favours a ?hierarchical scheme@ of develo#%ent$ which #ortraits an extre%e
version of the neo(arwinist view of genes directly #rescribing develo#%ental
#rocesses in turn s#ecifying %or#hology in a straightforward way! "n it both
%or#hological evolution and develo#%ent are reduced to #urely genetic #roble%s&
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
develo#%ent to a se2uence of gene ex#ression$ and evolution to a change in gene
fre2uencies!
4owever$ this sche%e #resents several #roble%s )*++*0! <irst$ such an ?o#en loo#@
syste% would be extre%ely unstable against the rando% disturbances that acco%#any
nor%al develo#%ent! Decond$ the effect of genes on %or#hology is %ostly indirect& they
code for %olecules which either regulate the ex#ression of other genes or confer
#ro#erties on cells )e!g! cell division rates$ a#o#tosis$ differentiation ti%ing or
cytoskeletal #ro#erties0$ which then construct organs and structures largely in
accordance with #hysico(che%ical laws! Thus$ the co%#lexity and nonlinearity of the
genoty#e(#henoty#e %a##ing increases at higher levels of interaction! This ex#lains
why we do not find a one(to(one corres#ondence between a given %or#hological trait
and a given region of NA )*+,90& due to the highly context(sensitive character of gene
ex#ression )de#endent on the stage and the local environ%ent0$ si%ilar genetic changes
%ay yield different %or#hological effects$ and the other way round! "n fact$ at the
%or#hogenetic level$ genetic %utations %ay not even be ex#ressed$ as develo#%ental
interactions have #ro#erties that e%erge fro% the dyna%ics of the syste% and are not
encoded in the geno%e )Bster C Alberch *+,=$ Bster et al! *+,,$ Alberch *++*0!
Therefore$ #henoty#ic diversity is not so %uch the #roduct of new genes as of
#er%utations in the context )i!e! the ti%ing and location of ex#ression0 of existing
genes! The evolutionary conse2uences of this asy%%etry are obvious& there are
2ualitative differences between %odes of evolution at the genetic and e#igenetic levels$
and therefore there is often no direct corres#ondence between genetic and
%or#hological divergence )*+,90!
"n order to ca#ture the #ro#erties of develo#%ental syste%s and their evolutionary
conse2uences$ Alberch #ro#oses an alternative view of develo#%ent which he calls the
?cyclical/feedback scheme@! According to it$ develo#%ental #rocesses are divided in
three interacting levels )*+,=a$ #! 9=-0& interactions a%ong genes within a highly
structured geno%e$ #rotein and enzy%e ones generating cell #ro#erties involved in
%or#hogenesis$ and tissue interactions! As Aaddington$ Alberch considered that these
regulatory interactions s#ecify the e#igenetics according to which #henoty#es are well
buffered syste%s with res#ect to both genetic and environ%ental #erturbations during
ontogeny )*+,-0! Therefore$ in the cyclical view of develo#%ent ?genes are :ust one
ste# in the chain of interactions$ gene ex#ression is both the cause and the effect of a
%or#hogenetic #rocess@ )*++*$ #! .0!
Thus$ Alberch>s %essage is that the %or#hogenetic level of descri#tion deserves s#ecial
attention in Evoevo because it vastly governs both the dyna%ics of %or#hogenesis
and the a##earance of novelty! <or Alberch it is at this level where the range of #ossible
variation for evolution is established$ and it introduces a deter%inistic factor in the
#roduction of variation )which was largely left to rando% %utations0 that allows so%e
for% of #redictability$ in so far as the variability of a given trait )its #otential for
#roducing variations0 can be established!
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
3. !evelopmental dynamics" from heterochrony to construction rules
es#ite the efforts of workers like de Eeer$ Aaddington or Dch%alhausen$ the role of
develo#%ent was absent fro% standard evolutionary theory since the decline of the
reca#itulation theory! 4owever$ it received a renewed interest by the scientific
co%%unity after the #ublication of 6ould3s #ntogeny and $hylogeny )*+110! Two years
later$ Alberch and co(workers #ublished together with 6ould a celebrated #a#er where
they #resented a 2uantitative %ethod for describing the relationshi# between
heterochronic changes and #hyletic trends )Alberch et al! *+1+0! The ai% was to
i%#rove 6ould>s 5clock %odel5$ which was essentially 2ualitative and 5static5! Thus$ in
the first #lace$ Alberch and co(workers 2uantified 6ould5s %odel$ %odelling
%odifications in the develo#%ental #rocesses that #roduce relative changes in size and
sha#e! "n the second #lace$ they gave a dyna%ic ex#ression of heterochrony$ defined in
ter%s of shifts in s#ecific #rocesses such as onset$ cessation$ or rate of growth$ rather
than of end results! This a##roach is followed in later #ublications$ where we find
e%#irical work on heterochrony in Bolitoglossa occidentalis )Alberch C Alberch *+,-0
This treat%ent of heterochrony )and their focus on #atterns of relative growth0 was very
influential and acted to define the way the conce#t is used in current Evoevo )D%ith
=--*$ =--=0! 4owever$ after *+,/ Alberch found that this a##roach had serious
#roble%s$ and revised his views in accordance to his %ore dyna%ical and %echanistic
conce#tion of develo#%ent )*+,/a$ Alberch C Elanco *++.0! Alberch o##oses two
conce#tions of ontogeny and #hylogeny& a kinematic one% which only classifies and
co%#ares develo#%ental stages$ and a dynamic one based on an understanding of the
underlying %echanis%s of develo#%ent and the co%#arison of develo#%ental events
)*+,/a$ Bster et al! *+,,0! The conce#tion of develo#%ent as a se2uence of
discontinuous %or#hological stages conserved throughout evolution and the associated
%echanis% of heterochrony takes roots in a 4aeckelian a##roach! Eut$ recovering the
criti2ue %ade by von Eaer to the law of #arallelis% of ;eckel(Derres$ Alberch re%inds
that co%#arative e%bryology does not show any linear reca#itulation$ and concludes
that all the inter%ediate stages #ostulated by heterochronic %odels are %eaningless
)*+,/a$ #! /*0! "nstead$ changes between two related %or#hologies ?%ust be searched
for in ter%s of changes in the develo#%ental rules of interaction or initial conditions$
rather than in inter%ediate ontogenetic stages@ )#! /*0! 4e uses several exa%#les$
including the ex#eri%ents of #revious #a#ers )Alberch C 6ale *+,9$ *+,/0 to ?illustrate
how evolutionary %odels based on heterochrony can be e%bedded within a %ore
%echanistic and dyna%ic fra%ework@ )*+,/a$ #! //0! 4e had to say then that the work
done in *+1+ was ?no longer valid@ )#! //0$ because this #ers#ective did not fit well
with his current views of develo#%ent as a dyna%ical syste% governed by a set of
?construction rules@ that are able to generate a global #attern!
This new conce#tualization of develo#%ent has i%#ortant conse2uences on the
definition of ho%ology$ a hot 2uestion in current evolutionary biology! The distinction
between develo#%ental stages and develo#%ental events leads Alberch to ?cha%#ion
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
the view that %any of the difficulties in establishing ho%ologies could be avoided or
resolved by basing co%#arisons between ele%ents on the develo#%ental #rocesses
which created the%$ rather than on their final geo%etric for%!@ )Bster et al! *+,,$ #!
,110! The best e%#irical exa%#le of this a##roach is the study of the %or#hogenesis of
the vertebrate li%b! The early stages of the skeletal #atterning were ex#lained by a
%echanistic %odel of e%bryonic branching and seg%entation in initial chondrogenesis!
"n this %odel$ the loss of a digit %ay result fro% the failure of a branching bifurcation$
and then$ it is not sensible to ask 5which5 digit was lost$ since it is the basic se2uence
what has been altered! Thus they are not the %or#hological ele%ents but the
%or#hogenetic #rocesses what can be co%#ared )Bster et al! *+,,0!
"n order to ca#ture this new view on develo#%ental evolution$ Alberch develo#s the
notion of construction rules% which crystallized in #ublications on the %or#hogenesis of
teratologies and tetra#od li%bs! The co%#arative analysis and the ex#eri%ental
%ani#ulation show how these rules deter%ine how a li%b can be %odified by evolution&
An understanding of these$ very often si%#le$ rules can #rovide valuable insights
into the a##arent co%#lexity of %any develo#%ental #rocesses and$ further%ore$
allow us to deter%ine the relationshi#s a%ong different #henoty#es$ since the set of
#ossible #henoty#ic transfor%ations will be constrained by the generative
#otentialities of the %or#hogenetic rules involved in the #rocess! )*+,=$ #! 9=*0
evelo#%ent reveals as a dyna%ical syste% governed by certain rules or constraints
which re%ain stable during long #eriods of ti%e$ but #er%it certain range of variation
through the alteration of certain )genetic and non(genetic0 #ara%eters! These rules arise
fro% the interaction of different 5resources5 at different levels and direct or channel the
#ossible evolution of the syste%!
3.3. The formal properties of developmental systems and the role of
mathematical biology
"n order to ex#lain the %echanics of develo#%ental #rogra%s$ Alberch used the
conce#tual and %athe%atical tools develo#ed in the fra%ework of dyna%ical syste%
theory! evelo#%ental #rocesses are viewed as ?co%#lex dyna%ical syste%s$ where a
s%all set of si%#le rules of cellular and #hysico(che%ical interaction can interact to
generate a co%#lex %or#hology@ )Bster C Alberch *+,=$ #! 7//0!
These rules of construction$ #robably ins#ired in the conce#tual fra%ework of cellular
auto%ata$ are for%ally ca#tured as develo#%ental #ara%eters! Alberch #ostulates that
develo#%ental #rocesses$ as any #attern(generating syste%$ have an associated
#ara%eter s#ace that ca#tures the global #ro#erties of the syste%$ defining its
?evolutionary #otential@ or evolvability&
This a##roach views for% and #attern e%erging as the co%bination of a set of
s#ecific #attern(generating algorith%s with a set of initial boundary conditions!
iversification occurs as the result of changes in the develo#%ental #ara%eters of
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
the algorith% )F@#rogra%@ co%#osed of a set of local rules of bioche%ical$ cell(
cell$ or tissue interactions0! The basic structure of a #attern for%ation %odel is to
2uantitatively state so%e local rules of interaction that are able to generate a global
#attern )*+,/a$ #! /-0
The effects of genetic or environ%ental alterations in the basic develo#%ental #ro#erties
are %athe%atically abstracted as #ara%eter #erturbations in a dyna%ical syste% )*+,=$
#! 9=90! Thus$ ?%or#hological diversity is generated by #erturbations in #ara%eter
values )such as rates of diffusion$ %itotic rate$ cell adhesion$ etc!0 while the structure of
the interactions a%ong the co%#onents re%ains constant@ )*+,+$ #! =10! "n order to
visualize all #ossible #athways of transfor%ation a%ong #henoty#es that so%e s#ecific
rules of construction generate$ Alberch i%#orted a tool fro% dyna%ical syste%s theory&
transformational diagrams )*++*0! Each s#ecies or trait will have a uni2ue
transfor%ation diagra% de#endent on its #osition in the #ara%eter s#ace$ and s%ooth
#erturbations of the #ara%eters can result in only certain given #henoty#es! These
#ro#erties of develo#%ental syste%s are illustrated by teratologies$ which are not only
generated in an organized and discrete way$ but also exhibit general transfor%ational
rules! Like in the case of the li%b of sala%anders$ the #erturbation of develo#%ental
#aths through a genetic %utation or ex#eri%ental %ani#ulation can only #roduce a
li%ited set of transfor%ations!
Alberch considers that the stabilities and bifurcations of Aaddington5s e#igenetic
landsca#e can be now for%alized with the language and %athe%atical tools develo#ed
in dyna%ical syste% theory& #henoty#ic stabilities are treated as dynamical attractors$
regions in #ara%eter s#ace where s%all #erturbations do not disru#t the basic
organization! "n the sa%e way$ ;Gller and Aagner state that ?ontogenies can be
understood as syste%s of te%#orary e2uilibria or steady states between develo#%ental
entities@ )*++*$ #! =7+0! Bifurcations are develo#%ental thresholds )such as critical cell
nu%ber or inductive or s#atial relationshi#s0 with the o##osite result! The way to
ex#lain how %echanistic rules can constrain #rocesses of %or#hological evolution has
to do with the #resence of %athe%atical bifurcations and how it ?i%#lies discontinuity
and directionality in #hyletic transfor%ations@ )Bster C Alberch *+,=$ #! 7//0! Thus$
develo#%ental thresholds ex#lain the nonlinear character of develo#%ent and the
discontinuity of evolution outlined by the theory of #unctuated e2uilibriu% )Eldredge C
6ould *+1=0& %odifications that go beyond such thresholds can cause nonlinear effects!
3.&. The physical properties of developmental systems and the role of
experimental embryology
Theoretical research ca#tures the structural #ro#erties of develo#%ental syste%s$ but the
study of the %echanical and che%ical laws to which the %or#hogenetic #rocesses are
sub:ected is necessary )Bster et al! *+,,0! Alberch was not :ust a theoretical biologist$
but a brilliant ex#eri%ental e%bryologist! The #a#ers #ublished in collaboration with
E%ily 6ale since *+,9 illustrate this work )Alberch C 6ale *+,/$ *+,.H Alberch *+,.H
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
Alberch et al! *+,.0! 4ere the influence of #erturbations of so%e develo#%ental
#ara%eters in the generation of new for%s is studied ex#eri%entally! The authors
co%#ared the results of treating the li%b buds of a frog and a sala%ander with
colchicine$ a %itotic inhibitor! This treat%ent results in various abnor%al %or#hologies
such as li%bs of s%aller size that have lost so%e skeletal ele%ents! Thus they can
contrast the hy#othesis that the digital #attern is affected by reduction in the nu%ber of
%esenchy%al cells! Ihanges in #attern for%ation take #lace when the size and the
nu%ber of cells of the li%b bud are reduced under a critical value$ an ex#eri%ental fact
consistent with the %athe%atical %odels studied with Bster )Bster et al! *+,,0! Eut the
%ain result is that the %alfor%ations #roduced by colchicine are not chaotic$ but they
exhibit a high degree of order! ;ost of the #atterns of diversity of digital %or#hology in
a%#hibians can thus be ex#lained as a reflection of develo#%ental #ro#erties )Alberch
C 6ale *+,/0!
The conse2uences of the ex#eri%ental %ani#ulation of develo#%ental syste%s are
twofold! "n the first #lace$ the ex#eri%entally generated #attern of variation can be
co%#ared with the #atterns of natural variation$ allowing #hylogenetic inferences and
to trace #ossible evolutionary #athways! <or exa%#le$ sala%anders see% to develo#
their li%bs in a way very different fro% the rest of tetra#ods because the se2uence of
digit for%ation see%s to be inverted )Dhubin C Alberch *+,.$ Alberch C 6ale *+,90!
igit reduction is a #heno%enon that has taken #lace several ti%es in an inde#endent
way in a%#hibian evolution! <rogs usually loose their %ost internal digit )#reaxial0
whereas sala%anders the %ost external one )#ostaxial0! The interesting result obtained
in this co%#arative study is that there is a #arallelis% between ex#eri%entally generated
#atterns and the evolutionary trends towards digit reduction observed in wild ani%als
)Alberch C 6ale *+,/0! At the sa%e ti%e$ the bounded variability generated in the
laboratory reveals a funda%ental #ro#erty of develo#%ental syste%s! Although not all
induced %alfor%ations #roduce for%s observed in nature$ these non(natural
%or#hologies are also li%ited by the syste%3s %or#hogenetic #ro#erties$ that is to say$
they are the #roduct of the iteration of rules! Dhubin and Alberch )*+,.0 %aintain that
during the evolutionary history of vertebrates these basic %or#hogenetic rules have not
changed$ only the interaction #ara%eters! Juantitative variations of these #ara%eters
%ay #roduce 2ualitative alterations such as changes in the branching and seg%entation
se2uences$ but since the rules of interaction re%ain the sa%e$ we will never obtain
2ualitatively new for%s in the laboratory$ but atavis%s$ deletions$ or du#lications of
already existing features )*++*0! "f we are dealing with an evolutionarily fixed
develo#%ental syste% we are only able to ex#lore its #otentialities$ and %ostly
reiterating for%s that had already been realized during evolution )*++*0!
!. The role of developmental constraints
Alberch dee#ly ex#lored the notion of ?develo#%ental constraint@ of the Spandrels
#a#er )6ould C Lewontin *+1+0$ both in a theoretical )*+,=$ *+,.$ *+,+0 and in an
ex#eri%ental way )Alberch C 6ale *+,9$ *+,/0! The notion of constraint co%es to
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
biology fro% other fields dealing with co%#lex syste%s$ such as #hysics or engineering$
where it is often understood as a li%itation of variation of so%e kind$ so%eti%es with a
creative or enabling character! "n fact$ this sense of a li%itation of variety at one level
co%bined with the e%ergence of #ossibilities at a higher level is the one Kacob uses in
his also influential #a#er )*+110! 4owever$ when 6ould C Lewontin )*+1+0 introduce in
the literature the notion of 5develo#%ental constraint5$ the ai% is to criticise the view of
evolutionary ada#tation as #erfect o#ti%ization through natural selection$ and there the
notion of constraint does not refer to how levels integrate in a co%#lex syste% but to a
li%itation of the #ower of natural selection considered as an ada#tive force&
This #rogra%%e regards natural selection as so #owerful and the constraints upon
it so few that direct #roduction of ada#tation through its o#eration beco%es the
#ri%ary cause of nearly all organic for%$ function$ and behavior! )6ould C
Lewontin *+1+! Bur e%#hasis0
Dince then$ the notion has had to struggle between these so%ehow difficult to conciliate
two senses of li%itation )of natural selection3s #ower0 and organization )of a co%#lex
%or#hology through an evolved develo#%ental dyna%ics0$ and there have been
atte%#ts to clarify the% )see A%undson *++7$ Dchwenk *++/$ 6arcia(Azkonobieta
=--/0! A%undson )*++70 distinguished between constraints on ada#tation Lwhich
would #revent certain changes however environ%ental #ressures workedMand
constraints on for% Lwhich would be indifferent to ada#tation and only had to do with
#ossible %or#hologies! <or hi%$ the strength of each of the two %eanings was %ore
evident if we look at the% as being #art of different research #rogra%s or traditions$
na%ely that of neo(arwinis% )externalis%0 in the first case and that of
'aturphilosophie )internalis%0 in the second!
Alberch distinguished a range of constraints that act at various levels and have different
degrees of generality$ but he was interested on constraint at a #articular level& the
cellular and tissue as#ects of #attern for%ation and %or#hogenesis )*+,/b0! These are
constraints involving %or#hogenetic construction rules$ res#onsible of the fact that
when organis%s de#art fro% their nor%al develo#%ent$ they do so in only a li%ited
nu%ber of ways!
Alberch5s definition of develo#%ental constraint fits in the constraint on for%$ and thus
re%ains organizational&
"f the raw %aterial u#on which selection o#erates is generated by effectively
rando% and boundless variation$ anything will be #ossible! Therefore it cannot be
#redicted$ exce#t by using o#ti%ality argu%ents and assu%ing that$ if so%e
#henoty#e would be o#ti%al$ the a##ro#riate genoty#es will a##ear! )N0 An
alternative a##roach to this organis%(environ%ent dichoto%y consists in focusing
on the internal organization of the organis% and atte%#ting to characterize the
constraints operating on the system' )*+,=$ #!9*7! Bur e%#hasis0
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
A develo#%ental constraint is a bias in the ex#ression of #henoty#ic variability due
to the nature of the interactions that characterize develo#%ental syste%s! )*++-$ #!
*/0
<or hi%$ constraints reflect the intrinsic abilities of develo#%ental syste%s to #roduce
so%e for%s and not others! This fra%ework has certain conse2uences about the
generation of the variation re2uired for evolution to #roceed$ as ?the inter#lay between
genetic indeter%inacy and resilient develo#%ental rules suggests the likely recurrence
of si%ilar %or#hological variation and the generation of a biased subset of #henoty#es
u#on which natural selection or #o#ulation stochastic factors can o#erate@ )Alberch C
6ale *+,/$ ##! *+(=-0! Therefore$ in this view e#igenetic interactions do not constrain
natural selection$ but how genetic %utations are ex#ressed at the %or#hological level!
Thus$ the conce#t of develo#%ental constraint illustrates the need of an elucidation of
the develo#%ental structure of the syste% in deter%ining the nature of the
develo#%ental constraint on #henoty#ic variation! <ro% this #ers#ective$ constraints
are not only restrictions for certain conceivable novelty in evolutionH they #rovide
#ossible directions for changes and innovation that we actually find in the biological
real%! "n other words$ constraints are not only i%#ortant to tri% ada#tationist o#ti%ality
thinkingH they are a funda%ental tool for the evolutionist who wants to search both for
the source fro% which certain innovations e%erged and #ossible directions for future
evolutionary change! An illustrative exa%#le co%es fro% Alberch5s study of the
evolution of %eta%or#hosis )*+,10! "n this #a#er$ he hy#othesizes that the #arallelis% in
the evolution of %eta%or#hic #atterns in insects and a%#hibians suggests a co%%on
develo#%ental constraint& the irreversibility of develo#%ent i%#oses a general
constraint that sha#es #hylogenetic #atterns of %eta%or#hosis$ since such distant #hyla
circu%vent it with the si%ilar evolution of redundant syste%s! Thus$ the general
constraint of the irreversibility of develo#%ent does not li%it the generation of
variation$ but %akes #ossible the a##earance of certain new develo#%ental 5strategies5 in
the generation of %or#hological novelty!
Resnik3s )*++/0 distinction between activeO#assive develo#%ental #atterns or
constraints can hel# us to clarify this view! The ?rules of construction@$ as defined
above$ guide the #rocess of for% construction! These rules are then active
developmental constraints" 'e#igenetic interactions drastically constrain the universe of
#ossible %or#hological novelties and impose directionality in %or#hological
transfor%ations though #hylogeny' )*+,-$ #! ./7! Bur e%#hasis0! The dyna%ical
#ro#erty of the directionality in the transfor%ations a%ong steady states has a s#ecial
relevance because it gives us a cue to understand the 5active5 or 5#ositive5 %eaning of
constraint that Alberch has in %ind )also defended by 6ould *+,+0! yna%ical syste%s
tend to fall in their attractors following so%e s#ecified directions! Do Alberch3s rules or
active develo#%ental constraints deter%ine not only the actual for% but the set of all
#ossible #henoty#ic transfor%ations in virtue of its generative #otentialities! They are
active developmental constraints on evolution in Resnik3s sense! Transfor%ational
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
diagra%s show the #otential evolutionary transfor%ations of #henoty#es #redicting the
%ost #robable one in absence of external forces! They can be used as a 5null hy#othesis5
of evolutionary transfor%ations$ since Delection only can drive #henoty#es along the
internally s#ecified directions! Thus$ in Alberch3s view$ the dyna%ical #ro#erties of
develo#%ental #atterns li%it the variability #roducing clusters or attractors in the
#henoty#ic of #ara%eter s#ace$ but at the sa%e ti%e$ #rovide #otential directions to
evolutionary change!
The #ositive %eaning of develo#%ental constraints is s#ecially illustrated in Alberch5s
ideas on evolvability$ now a core conce#t in Evoevo )Aagner C Altenberg *++.$
8irschner C 6erhart *++,0! After awkins )*+,+0 established the ter%$ he dedicated a
#a#er to the different abilities of develo#%ental syste%s to evolve )*++*0! The
#ro#erties of develo#%ental syste%s )stability and variability based on develo#%ental
thresholds0 define their evolvabilityH the ability to evolve re2uires that the syste% has
stability against #erturbations$ but not so %uch as to be i%%une to absorb change and
variation! These very general #ro#erties of develo#%ental syste%s lead Alberch to
#ro#ose a new level of selection&
there %ust have been selection a%ong #attern generating syste%s favouring the
ones that exhibit the ade2uate balance between stability and #otentiality to generate
sufficient #henoty#ic variability! "t %ust be e%#hasized that we are dealing here
with a new level of selection$ one that it does not act on the #henoty#e nor on the
genoty#e$ but rather on the e%ergent #ro#erties of develo#%ental syste%s )*++*$ #!
*-0
Alberch #resents the generation of for% as a #rocess governed by certain generative
rules or constraints that both #rovide stability and #ro%ote change in certain directions!
Eut$ can this view ex#lain the creative ca#acity of evolution$ for instance$ in the origin
of new body #lansP
The answer to this 2uestion is$ in general$ #ositive$ since it see%s that to talk about
creativity we need a reference to a s#ecific generative syste%$ or to so%e set of
constraints! "n this sense$ Alberch see%s to have thought that the eli%ination of all
constraints and their substitution by rando% #rocesses will su##ress creativity!
4owever$ the conservation of sets of interaction rules within ontogenetic ty#es
constrains the range of creativity of develo#%ental syste%s! The ex#loration of the
s#ace generated by the develo#%ental rules can bring %ore or less creative results$ for
exa%#le$ a co%#lex #athway in the transfor%ational diagra%$ with a lot of transitions$
or the introduction of %odulation in %usic! Eut the %echanis% that can #roduce really
creative for%s is the transfor%ation of the generative s#ace by the change or the
eli%ination of so%e of the rules! <or instance$ this is how the su##ression of the axio%
of #arallels resulted in the discovery of the non(Euclidean geo%etries! Alberch3s
#resents the Ia%brian ex#losion$ one of the %ayor creative events in evolution$ as an
exa%#le of this for% of creativity$ with ?the invention of %ulticellularity$ seg%entation
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
or the se2uestration of the ger% line a##ear$ with hindsight$ to have been key
develo#%ental events that have s#eeded u# the evolutionary #roliferation of lineages@
)*++*$ #! +0! Alberch thinks that the Ia%brian was a #eriod of ex#eri%entation in rules
of cell(cell interaction$ rules that exhibited different for%(generating abilities as well as
distinct stability #ro#erties )*++*0! After it no 2ualitative new structural body #lans
see% to have a##eared and %or#hological variation looks reduced to variations within
the%es!
Accordingly$ it is i%#ortant to kee# in %ind that although Alberch insisted in
#redictability and deter%inis%$ this is only within the fra%ework of a given
develo#%ental syste%! Bne of the %ain criticis%s to the notion of develo#%ental
constraint is that constraints can be broken in the course of evolution! Dalazar(Iiudad
)=--/0 has #ro#osed to overco%e the divide between the standard neo(arwinian
a##roach and the ?Ionstraint school@ by studying the variational properties of different
ty#es of develo#%ent! This idea is congenial to Alberch>s thought! "f evolution can
switch develo#%ental ty#es$ then for%s of history and contingency can %aybe also
taken into account$ because then a ty#e of develo#%ent does not constrain evolution in a
deter%inistic way$ since the syste% can always switch into another ty#e! Qet$ each
develo#%ental ty#e can be said to be constrained in the sense that it generates a given
range of #atterns!
Alberch was interested in the e#iste%ology of creative #rocess as well$ which he
considered to be guided by certain internal rules! <or instance$ the are so%e basic rules
or #heno%enological #rinci#les in the creation of %ythological creatures$ in the sa%e
s#irit of the rules involved in the construction of tetra#od li%bs& a0 distortion of size
andOor for% of existing #arts$ b0 %ulti#lication or su##ression of an existing #art$ c0
reco%bination of #arts that eventually have suffered so%e of the #recedent
transfor%ations! <or hi%$ these universal %ythical for%s$ like natural for%s$ are not
%ere historical contingencies$ but reflect a dee# structure of %ind which is dyna%ical!
As the ro%antic #hiloso#hers and #oets that he cites$ he thinks that the %ental #rocess
of creation of a new for% is a %or#hogenetic #rocess that de#ends on brain dyna%ics
)*++,0!
". #onclusions: Towards an interative bioloy
Aagner and co(workers )=---0 distinguish three #hases in the constitution of Evoevo
as a disci#line! They suggest that the ahle% conference )Eonner *+,=0$ where Alberch
was one of the central figures$ %arks the end of the ro%antic stage! The enthusiastic
#hase started later with the discovery of ho%ologous genes in !rosophila and other
ani%als! Ae will be nowadays in the #rocess of the constitution of the academic stage$
where develo#%ental %echanis%s and evolutionary #rocesses should be integrated! "t
see%s to be an overall agree%ent that the %ain challenge to current Evoevo is the
integration of different theoretical fra%eworks in establishing the link between
develo#%ent and evolution! The #roble% is that %any authors see develo#%ental
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
evolutionary genetics as the new 5synthesizer5 of evolutionary biology )6ilbert =--9$
A%undson =--/0! "n this view$ unity of type is revealed by 4ox genes$ whereas
ada#tations to conditions of existence are assi%ilated by in genetic networks and filtered
by natural selection! 4owever$ we think that work of an Alberchian style$ involving
%or#hological research to establish #atterns of variation and ex#laining variability at
the e#igenetic level is funda%ental for the integration of the diverse research #rogra%s
converging in Evoevo )If! ;Gller C New%an =--/0! That is why$ to conclude$ we
would like to su%%arize so%e of the #hiloso#hical issues ste%%ing fro% Alberch>s
work that #oint towards a true integrative biology&
(nternalism.) "n Alberch>s #a#ers one #erceives a resistance to a given way of thinking
about biology )in broad ter%s$ what 6ould and Lewontin )*+1+0 called ada#tationis%0$
and an a##eal to a different view of organis%s and their #rocesses for which he used
different designations$ like 'internalis%'&
" focus on the internal rules that control the a##earance of %or#hological variation$
on the %echanistic basis of such rules and on the evolutionary conse2uences of this
internally deter%ined order )*+,+$ #! =,0!
Ae think that Alberch>s target of criticis% is not the hy#othesis of natural selection by
itself$ but the a##roach that finds that ada#tation to an external situation )the
environ%ent0 is the only source of living organization! Eiology as he conceived it would
#ay a closer attention to the %echanis%s and regularities ste%%ing fro% an organic
5internal order5$ but this belief does not lead hi% to dis%iss the role of natural selection&
?Brder in evolution is a co%bination of deter%inistic agents at two levels& internally
generated order based on the internal dyna%ics of develo#%ent$ and natural selection$
de#endent on the #ro#erties of organis%(environ%ent interaction!@ )*+,+$ #! 7.0! As a
%atter of fact$ both sources of order are not inde#endent 5filters5! Bn the contrary$ in
Alberch5s view$ the evolutionary #rocess is dee#ly interactive! Ahen acting u#on a
#articular #henoty#e$ selection indirectly affects the develo#%ental syste% and this
creates a series of contingent history(de#endent evolutionary #otentialities! Delection
o#erates within a set of internally defined #ossible transfor%ations and when it
deter%ines the #robability of survival of each of the%$ it indirectly affects the
%or#hogenetic #ro#erties of the s#ecies! "t is only because of the stage in which this
field of study still is that he #refers to #ush the internalist a##roach&
At this stage of develo#%ent of the field$ it is %ore useful to study the organization
and evolution of for% fro% the internalist #ers#ective! This a##roach$ based on an
understanding of develo#%ent$ naturally incor#orates the #heno%ena of regulation$
integration and constraint that are inherent to the conce#t of organis% )i!e! for%0
and which are absent in the conce#tual fra%ework #rovided by the externalist
a##roach which tends to view the #henoty#e as a static entity! )*+,+$ #! 7,0
Bne has to envision that within an integrated fra%ework all as#ects of evolution %ust
be co%#le%entary& the %or#hological accounting for the generation of for%$ the genetic
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
standing for %olecular details and the interactive understanding )between for%s and
with the environ%ent0 to integrate functions!
Form and function.) Related to the need of bringing forth the internalist #ers#ective is
the vindication of a 'theory of for%' based on the global #ro#erties of the network of
interactions that characterize develo#%ent&
Evolutionary biology needs a theory of for% RNS The conce#ts of regulation and
feedback that characterize develo#%ental syste%s are cons#icuously absent fro%
existing theoretical sche%es )*+,+$ #! 9+0!
Alberch5s contribution is in our view relevant to #ro%ote the inclusion of studies centred
in the interactive dyna%ics of #rocesses of for% generation within the theoretical
fra%ework of Evoevo! "n fact$ his last works intended a general theory of for%$ not
:ust in biology$ but also in technology and art! There he started fro% the study of #attern
generation in co%#lex syste%s to a##roach the #roble% of for%s without functions$ i!e!
for%s not attending to external functional de%ands )*++,0! ;onsters were definitely for
hi% an exa%#le of for%s without function$ and so were %achines without functions$
?%onstrous %achines@ as he called the%!
4owever$ Alberch does not restrict the %eaning of function to the external %eaning of
natural selection! Recovering the Iuverian tradition )see also Dchwenk =--*$ Reiss
=--/0$ he defends an internal a##roach to function that accounts for the organis%al
integration of for%s$ defining functional constraints as those ?i%#osed by functional
interactions a%ong different #arts of the organis%@ )*+,*$ #! ,70! Do Alberch foresaw
an a##roach to functions that started with both the generative and interactive #ro#erties
of #atterns$ and not with the ada#tive advantages assigned to se#arate #arts! <ro% this
view$ the #rocess of #roducing integrated and organized #henoty#es beco%es a three
ste# one of a0 generation of for%s )%or#hogenesis0H b0 integration of those for%s
through functional interactionsH and c0 natural selection and other contingent historical
#rocesses! Thus$ new %or#hologies are first #roduced$ and then these novelties have to
be integrated in the rest of the organis% interacting with the environ%ent )*+,=b0!
*echanistic typology.) Alberch5s distinction between the kine%atics )descri#tion0 and
the dyna%ics )ex#lanation0 of evolutionary change reveals an organized synthesis of
#hylogenetic research& the co%#arative evolutionary a##roach to #hylogenetic #atterns
and the %echanistic analysis of develo#%ental #rocesses! Thus$ %or#hological
disci#lines )co%#arative anato%y$ taxono%y$ syste%atics and #alaeontology0 are
necessary to characterize the #atterns of variation$ while ex#eri%ental e%bryology and
%athe%atical biology$ in #er%anent dialogue with evolutionary %or#hology$ will hel#
to discover the %echanis%s res#onsible of such variation!
The exclusion of %or#hology in conte%#orary Evoevo can be related to its
relationshi#s with ty#ology )Love =--90! The substitution of ty#ological thinking by
#o#ulational thinking was seen by ;ayr as the great con2uest of arwin$ and any kind
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
of ty#ology was censured as narrowly linked to essentialis% and idealis%$ the big
obstacles to evolutionis%! 4owever$ if we take into account Alberch5s work$ the
#roble% does not lie in the very ty#ology$ which accounts for the funda%ental
%or#hological di%ension of evolution$ but in the traditional statical and %erely
descri#tive a##roach to for%&
)T0his ty#ological and static a##roach is not o##osed to evolution or even to
natural selection! )N0 it does not draw fro% evolutionary %echanis%s based on
environ%entally defined selection and rando% %utation! The 2uest for a general
set of #rinci#les of for% is legiti%ate if we exchange the %eta#hysical conce#t of
the Bauplan for a %echanistic one based on #rinci#les of %or#hogenesis and
internal integration! )Dhubin C Alberch *+,.$ #! 9110
Thus$ Alberch5s work de%onstrates a for% of ty#ological thinking for%ulated in a
#urely %echanistic context$ disconnected fro% %eta#hysical references to i%%utable
essences )A%undson =--*$ Love =--90! As we have seen in his analysis of tetra#od
li%b$ this ty#e is not an ideal entity$ but the result of a historically conserved
develo#%ental #rocess$ non reducible to an exclusively genetic ex#lanation$ and which
serve to establish the range of #ossible variation u#on which selection acts!
+ontingency and determinism.) The study of variability obliges to a radical
e#iste%ological change in the do%ain of evolutionary theory because it introduces a
deter%inistic factor in the #roduction of variation as well as so%e for% of #redictability!
;or#hological coherence will be a source of deter%inis% absent fro% those accounts in
which the a##earance of novelty is considered to co%e only fro% rando% %utations&
An alternative a##roach to this organis%(environ%ent dichoto%y consists in
focusing on the internal organization of the organis% and atte%#ting to
characterize the constraints o#erating on the syste%! These constraints define the
#otential #athways of transfor%ation and i%#ose an additional deter%inistic
co%#onent on evolutionary #rocesses@! )*+,=$ #! 9*70
4owever$ Alberch does not defend an absolute but a ?relative@ deter%inis% )*+,*0! Ae
can say that throughout his work$ the e#igenetic level e%erges as a real% of
deter%inis% between two sources )#hysical and historical0 of uncertainty! <ro% the
#oint of view of develo#%ent$ Alberch #refers the ter% ?develo#%ental bias@ to that of
?develo#%ental #rogra%@$ because the latter has ?#erha#s a %ore deter%inistic
connotation than we intend here@! Although e%bryogenesis %ay a##ear deter%inistic on
a %acrosco#ic scale$ the cellular level #roceeds in a %ore stochastic fashion! ?"t is only
the aggregate behaviour that exhibits overall coordination!@ )Bster C Alberch *+,=$ #!
7770! The other source of un#redictability is the historical contingency co%ing fro% the
level of the interaction of develo#%ental constraints and natural selection& the %ost
#robable for%s in ter%s of their dyna%ic #ro#erties )stability or accessibility in the
transfor%ational diagra%0 can or cannot be #referred by selection! ;aybe an
Etxeberria, A., and L. Nuo de la Rosa. 2009. A World of Opportunity Witin !onstraint" #ere
Alber$%s Early E&o'de&o.( )n Pere Alberch. The Creative Trajectory of an Evo-devo Biologist,
edited by *. Rass+in',ut-an and .. *e Ren/i, 20122. 3alen$ia" #ubli$a$ions 4ni&ersitat de
3al5n$ia.
i%#robable for% can be the fittest in a deter%ined environ%ent! Rando%ness$
deter%inis% and contingency coexist in biological #rocesses! That is why ?we live in a
world of o##ortunity within constraint@ )*+,.$ #! ,0!
Ac$nowledements
This research received funding fro% grants "T(=/-(-1 fro% the Eas2ue 6overne%ent
and 4T;=--/(-=77+O<"DB fro% the D#anish ;inistry of Dcience and Education
);EI0 and <eder funds fro% the Euro#ean Tnion )AE0$ and a Predoctoral fellowshi#
fro% the Tniversidad Io%#lutense de ;adrid )E*+-U-70 )LNR0! Ae thank the initial
in#ut of To%as 6arcia Azkonobieta$ who contributed to a very early draft of this #a#er!
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