A	Developmental	Systems	Account	of	Human	Nature1 Karola	Stotz	(Macquarie)	and	Paul	Griffiths	(Sydney) Abstract It	is	now	widely	accepted	that	a	scientifically	credible	conception	of	human	nature	must	reject	the folkbiological	idea	of	a	fixed,	inner	essence	that	makes	us	human.	We	argue	here	that	to	understand	human nature	is	to	understand	the	plastic	process	of	human	development	and	the	diversity	it	produces.	Drawing	on the	framework	of	developmental	systems	theory	and	the	idea	of	developmental	niche	construction	we	argue that	human	nature	is	not	embodied	in	only	one	input	to	development,	such	as	the	genome,	and	that	it	should not	be	confined	to	universal	or	typical	human	characteristics.	Both	similarities	and	certain	classes	of differences	are	explained	by	a	human	developmental	system	that	reaches	well	out	into	the	'environment'. We	point	to	a	significant	overlap	between	our	account	and	the	'Life	History	Trait	Cluster'	account	of	Grant Ramsey,	and	defend	the	developmental	systems	account	against	the	accusation	that	trying	to	encompass developmental	plasticity	and	human	diversity	leads	to	an	unmanageably	complex	account	of	human	nature. 1.	Current	state	of	the	debate Human	Nature,	its	characteristics	and	its	causes,	is	one	of	the	oldest	and	most	contested	topics	of	inquiry. A	scientifically	credible	account	of	human	nature	must	assimilate	and	integrate	findings	from	the biological,	psychological,	and	social	sciences.	Contemporary	philosophical	work	on	human	nature	sets out	to	do	this,	but	it	also	tries	to	stay	in	touch	with	older	ideas	about	human	nature.	Almost	all	authors have	accepted	the	Darwinian	challenge	and	recognised	that	the	human	species	is	not	defined	by	a	fixed, inner	essence.	But	despite	this	rejection	of	essentialism	many	authors	remain	attached	to	the	idea	that human	nature	is	confined	to	the	left-hand	side	of	the	dicomies	between	nature	and	nurture,	innate	and acquired,	biology	and	culture.2 1	To	appear	as	Stotz,	Karola,	and	Paul	E.	Griffiths.	"A	Developmental	Systems	Account	of	Human	Nature." In	Why	We	Disagree	about	Human	Nature,	by	Tim	Lewens	and	Elizabeth	Hannon.	Oxford	&	New	York: Oxford	University	Press,	n.d. 2	Machery	2008;	Kronfeldner	In	Press;	but	see	Downes	and	Machery	2013	for	a	collection	of	different views;	Fuentes	et	al.	2010	for	a	collection	of	essays	providing	an	anthropological	challenge	to	a	unitary theory	of	the	human;	and	Lewens	2012	for	an	extremely	permissive,	if	not	eliminativist	notion	of	human nature. This	attachment	reflects	the	fact	that	enquiries	into	human	nature	start	from	an	everyday	('vernacular') idea	of	human	nature	and	try	to	honour	some	of	the	intuitions	associated	with	that	idea.	The	vernacular conception	of	human	nature	is	an	expression	of	an	implicit	'folk	theory'	of	biological	development	which has	at	its	heart	a	distinction	between	traits	that	come	from	'inside'	and	those	imposed	from	'outside'.	We and	our	collaborators	have	conducted	empirical	research	to	characterise	this	folk	theory	in	more	detail (Griffiths	2002;	Griffiths,	Machery,	and	Linquist	2009;	Linquist	et	al.	2011).	The	folk	theory	of	animal natures	is	an	instance	of	'psychological	essentialism'	(Medin	and	Ortony	1989;	see	also	Gelman	2003) and	the	essential,	inner	nature	of	an	animal	is	associated	with	traits	that	are	Fixed	in	development, Typical	of	the	species,	and	Teleological	–	the	animal	is	intended	to	have	this	trait.	When	this	folk	theory of	animal	natures	is	applied	to	humans	it	produces	the	vernacular	idea	of	human	nature.	We	describe	our 'three	factor'	model	and	related	psychological	research	in	Section	2. The	problem	with	the	vernacular	idea	of	human	nature	is	that	it	confounds	three	important	but essentially	independent	biological	properties.	A	trait	can	be	fixed	without	being	typical	or	having	a purpose,	it	can	be	typical	without	having	a	purpose	or	being	fixed,	and	it	can	have	a	purpose	without being	fixed	or	typical.	This	is	one	reason	why	so	many	developmental	biologists	and	psychologists	have rejected	a	simple	dichotomy	between	innate	and	acquired	characteristics	(Lehrman	1953;	Hinde	1968; Gottlieb	1970;	Bateson	1991). The	shortcomings	of	the	vernacular	idea	of	human	nature	are	similar	to the	shortcomings	of	the	pre-scientific	concept	of	heat.	Whether	an	object	feels	'hot'	depends	on	three physical	quantities	that	can	vary	independently	of	one	another	–	temperature,	quantity	of	heat	and conductivity.	Using	these	three,	more	precise	ideas	we	can	explain	what	people	are	responding	to	when they	say	something	is	hot,	but	the	original	idea	is	not	a	useful	construct	with	which	to	do	science. In	this	chapter	we	defend	a	view	of	human	nature	that	goes	beyond	the	vernacular	idea,	in	the	same	way that	the	physics	of	heat	went	beyond	the	phenomenological	notion	of	things	feeling	hot.	We	argue	that such	an	idea	must	fulfil	several	desiderata:	It	must	be	explanatory	and	not	merely	descriptive;	it	should make	human	nature	an	object	of	inquiry	in	the	human	sciences	(all	those	disciplines	that	take	the	human species	or	some	aspect	of	it	as	their	subject,	from	physiology	through	psychology	and	anthropology	to sociology);	a	science	of	human	nature	should	explain	the	folkbiological	features	traditionally	aligned	with the	idea	of	human	nature	and	why	they	won't	do	as	defining	features	of	human	nature;	and,	lastly,	our concept	of	human	nature	should	embrace	human	diversity,	plasticity	and	polymorphism,	because	these features	are	an	important	part	of	the	evolutionary	design	of	human	beings.	We	outline	these	desiderata in	more	detail	in	Section	3. We	will	argue	that	there	are	two	extant	theories	that	meet	these	requirements: Grant	Ramsey's	Life History	Trait	Cluster	(LTC)	account	(Ramsey	2013)	and	the	Developmental	Systems	(DS)	account	of human	nature	(Griffiths	2011).	In	Section	4	we	outline	the	basic	similarity	between	these	two,	namely that	both	are	grounded	in	human	developmental	biology.	Both	accounts	suggest	that	to	understand human	nature	is	to	understand	the	plastic	but	not	unstructured	process	of	human	development. While	we	are	in	agreement	with	much	of	Ramsey's	account,	in	section	5	we	draw	attention	to	some differences	between	the	two	accounts.	One	major	difference	is	that	our	account	focuses	more	strongly	on the	human	developmental	environment	as	a	critical	factor	in	human	nature.	Drawing	on	the	framework of	developmental	systems	theory	and	the	idea	of	developmental	niche	construction	we	argue	that	human nature	is	not	embodied	in	one	input	to	development,	such	as	the	genome.	The	patterns	of	similarity	and difference	amongst	human	beings	are	explained	by	a	human	developmental	system	that	reaches	well	out into	the	'environment'. We	also	emphasise	that	developmental	systems	theory	creates	a	dynamical,	process	perspective	on human	nature.	Human	nature	is	underpinned	by	a	range	of	mechanisms	of	extended	inheritance,	as	well as	genetic	inheritance,	and	the	life-course	of	any	individual	human	being	depends	upon	a	matrix	of exogenetic	developmental	factors	–	the	developmental	niche.	The	fundamental	unit	of	analysis	in	our theory	is	a	process	–	a	human	life-history	(Griffiths	and	Stotz	In	Press). 2.	The	folkbiological	idea	of	human	nature Our	account	of	the	folkbiological	conception	of	human	nature	builds	on	work	in	cognitive	anthropology and	child	psychology	that	identified	a	pattern	of	essentialistic	thinking	psychological	essentialism	about living	things	across	many	human	cultures	and	in	human	children	(Atran	1990;	Berlin	1992;	Medin and	Atran	1999;	Medin	and	Atran	2004;	Gelman	2003).	It	gains	additional	support	from	psychological research	on	the	'Genetic	Essentialism	Framework'	by	psychologist	Ilan	Dar-Nimrod	and	collaborators. Our	earlier	work	with	our	collaborators	constructed	a	'three-factor'	model	of	folkbiological	thought about	animal	natures;	provided	some	experimental	evidence	for	this	model;	and	showed	that	in contemporary	English	the	idea	of	'nature'	is	expressed	by	saying	things	are	'in	the	DNA'	(Griffiths, Machery,	and	Linquist	2009;	Linquist	et	al.	2011).	At	around	the	same	time,	Dar-Nimrod	and collaborators	set	out	to	study	lay	understandings	of	genetic	causation	and	documented	a	set	of	'genetic essentialist	biases'	that	correspond	closely	to	elements	of	the	three-factor	theory	of	animal	natures	(DarNimrod	and	Heine	2011a;	Dar-Nimrod	and	Heine	2011b;	Dar-Nimrod	and	Lisandrelli	2012;	Dar-Nimrod, Zuckerman,	and	Duberstein	2012;	Dar-Nimrod,	Zuckerman,	and	Duberstein	2014;	Cheung,	Dar-Nimrod, and	Gonsalkorale	2014). The	three-factor	model	proposes	that	there	is	a	folkbiological,	implicit	theory	of	development	in	which some	but	not	all	characteristics	of	animals	are	expressions	of	a	'nature'	inherited	from	their	parents	and which	makes	them	the	kind	of	animal	that	they	are	a	human,	a	chimp,	or	a	kangaroo.	Phenotypes	that stem	from	this	inner	nature	are	expected	to	have	three	characteristics:	Fixity,	Typicality	and	Teleology. Fixity	means	that	the	phenotype	is	hard	to	change	by	environmental	means.	Typicality	means	the phenotype	is	found	in	all	or	most	members	of	the	species	(or	of	some	natural	subset	such	as	a	sex	or	an age	group).	Teleology	means	that	the	phenotype	is	part	of	the	design	of	the	organism.	It	is	there	for	a reason,	and	organisms	that	lack	these	features	are	not	how	they	are	meant	to	be	(see	Section	3,	point	3 for	a	naturalistic	interpretation	of	Teleology).	In	Table	1	we	show	how	these	factors	line	up	with elements	of	the	Genetic	Essentialist	Framework	(GEF). The	GEF	suggests	that	genetic	attributions	for	various	traits,	conditions,	or	diseases	activate	four	specific psychological	processes,	or	genetic	essentialist	biases.	The	first	bias,	immutability/determinism,	is	that thinking	about	genetic	attributions	leads	people	to	view	relevant	outcomes	as	less	changeable	and predetermined.	To	the	extent	that	a	phenomenon	is	perceived	to	be	immutable,	it	will	be	perceived	to	be beyond	someone's	control.	Genetic	attributions	decrease	perceptions	of	control	over	relevant	outcomes (Dar-Nimrod,	Zuckerman,	and	Duberstein	2012;	Parrott	and	Smith	2014)	and	limit	the	perceived capability	of	other	means,	such	as	environmental	manipulations	or	individuals'	volition,	to	modify	the outcome	(Jayaratne	et	al.	2009).	The	second	genetic	essentialist	bias,	the	tendency	to	discount	additional causal	explanations	once	genetic	attributions	are	made	(termed	specific	etiology),	also	increases	the likelihood	that	people	will	disregard	alternative	casual	attributions	for	complex	phenomenon	(DarNimrod	and	Heine	2011a).	Whereas	the	first	two	genetic	essentialist	biases	focus	on	individuals,	the third,	homogeneity	/discreteness,	extends	attention	to	groups.	Essentialist	thinking	leads	a	person	to focus	on	the	central	identifying	features	that	are	common	to	all	group	members,	drawing	attention	away from	in-group	differentiating	features.	A	focus	on	commonality	should	lead	to	viewing	individual members	of	a	category	as	more	homogeneous	as	they	share	the	identifying	features,	which	may contribute	to	stereotyping	and	more	prejudiced	attitudes	toward	group	members	(Dar-Nimrod	and Heine	2011a).	The	final	genetic	essentialist	bias	is	naturalness,	that	is,	genetic	attributions	increase	the likelihood	that	a	relevant	outcome	is	perceived	as	a	natural	outcome.	There	is	long-standing	evidence	in psychology	that	viewing	an	outcome	as	natural	has	an	important	evaluative	component	attached	to	it. Genetic	Essentialist	Elements (Dar-Nimrod	and	Heine	2011a) Three-factor	model	of	animal	natures (Linquist	et	al.	2011) Immutable	and	determined:	thinking	about genetic	attributions	leads	people	to	view	relevant phenotypes	as	less	changeable	and	predetermined Fixity:	phenotypes	that	are	part	of	an	animal's nature	do	not	depend	on	the	particular environment	in	which	the	organism	is	raised	and are	hard	to	change	by	environmental manipulations Specific	etiology:	the	tendency	to	discount additional	causal	explanations	once	genetic attributions	are	made Traits	are	either	expression	of	the	animals	nature (and	are	expected	to	have	the	three	features)	or imposed	by	the	environment	(with	opposite expectations) Homologous	and	discrete:	leads	to	a	focus	on	the central	identifying	features	that	are	common	to	all group	members,	drawing	attention	away	from	ingroup	differentiating	features Typicality:	phenotypes	that	are	part	of	an	animal's nature	are	typical	of	the	entire	species	or	of	some natural	subset	such	as	males	or	juveniles Nature:	phenotypes	are	perceived	as	a	natural outcome	(with	positive	normative	associations) Teleology:	phenotypes	that	are	part	of	an	animal's nature	serve	some	purpose	(with	positive normative	associations) Table	1:	Comparison	between	the	Genetic	Essentialism	Framework	(GEF)	and	the	Three-Factor	Model The	vernacular	idea	of	human	nature	from	which	so	many	philosophical	analyses	start	is	simply	the application	of	this	form	of	essentialist	thinking	to	humans.	It	seeks	to	divide	human	characteristics	into those	imposed	by	the	environment	and	those	that	stem	from	an	inner	nature,	and	embodies	the assumption	that	the	three	characteristics	of	Fixity,	Typicality	and	Teleology	are	strongly	associated	with one	another	because	traits	that	stem	from	our	inner	nature	have	these	three	properties	and	traits imposed	by	the	environment	do	not. However,	this	intuitive	picture	of	biological	development	is	fundamentally	mistaken.	All	phenotypes	are produced	by	a	combination	of	genetic	and	environmental	factors,	and	in	many	cases	epigenetic	factors. The	patterns	of	interaction	between	these	factors	are	many	and	varied	and	do	not	conform	to	two distinct	patterns,	one	of	which	is	characteristic	of	traits	that	have	been	designed	by	natural	selection. Some	philosophers	have	conceded	this,	but	suggested	that	there	is	a	continuum,	with	evolved	traits clustered	at	one	end.	However,	in	our	view,	the	plausibility	of	this	idea	comes	not	from	reviewing	the evidence,	but	from	the	continued	influence	of	the	folkbiological	picture	(Griffiths	and	Machery	2008;	and see	Mameli	and	Bateson	2011;	Mameli	and	Bateson	2006). 3.	Desiderata	for	an	account	of	human	nature In	this	section	we	ask	which	desiderata	a	scientifically	credible	contemporary	conception	of	human nature	should	seek	to	fulfil. Kronfeldner	and	collaborators	have	distinguished	three	main	epistemic	roles for	the	concept	of	human	nature	(Kronfeldner,	Roughley,	and	Toepfer	2014;	Kronfeldner	In	Press).	The first	is	a	definitional	or	classificatory	role:	human	nature	defines	the	boundary	of	the	human	and determines	which	individuals	are	members	of	the	human	species.	The	second	is	a	descriptive	role:	the concept	collects	the	cluster	of	traits	characteristic	of	the	human	life	form.	This	can	be	seen	as	making human	nature	an	explanandum,	something	that	stands	in	need	of	explanation.	The	second	role	is therefore	complemented	by	a	third	role:	the	concept	as	an	explanans,	identifying	the	underlying mechanisms	or	factors	that	explain	why	humans	have	this	cluster	of	traits.	There	is	also	a	fourth, normative	role	for	the	concept	in	answering	the	question	of	what	a	'typical'	or	'proper'	human	ought	to be.	While	this	is	one	of	the	most	important	traditional	roles	of	the	concept	of	human	nature,	it	has	few supporters	in	philosophy	of	biology	(a	seminal	critique	in	this	field	is	Hull	1986). Philosophers	who	still try	to	use	human	nature	for	this	normative	purpose	do	not	derive	their	account	of	human	nature	from biology	(prominent	neo-Aristotelian	accounts	are	reviewed	in	Glackin	2015).	If	species	had	fixed,	typical, and	teleological	natures	in	the	way	that	folkbiology	supposes,	then	human	nature	could	fulfil	the	first three	of	the	roles	Kronfeldner	identifies,	and	perhaps	the	fourth.	But	since	there	are	no	such	natures,	a scientifically	credible	concept	of	human	nature	must	be	somewhat	revisionary.	It	will	give	people something	of	what	they	originally	wanted	from	a	concept	of	human	nature,	but	not	everything. We	believe	that	a	good	concept	of	human	nature	should	fulfil	the	following	desiderata: 1.	It	should	be	explanatory	and	not	merely	descriptive.	One	of	us	has	argued	elsewhere	that	a	purely descriptive	idea	of	human	nature	is	relatively	uncontroversial	(Griffiths	2011).	After	all,	there	are	a	range of	sciences	that	deal	with	humans,	and	many	of	these	sciences	are	successful,	which	implies	that	one	can abstract	away	from	the	particularities	of	individual	human	lives	to	discover	commonalities.	We	suggest that	in	addition	a	concept	of	human	nature	needs	to	address	what	causes	these	commonalities:	it	needs	to fulfil	an	explanatory	role. 2.	This	leads	to	our	second	desideratum:	a	useful	concept	should	make	Human	Nature	an	object	of inquiry	in	the	human	sciences,	namely,	the	sciences	that	deal	with	human	beings	as	a	kind.	For	example, physiology	tries	to	understand	functional	processes	in	the	human	body;	psychology	the	human	mind,	its underlying	processes	and	the	behavioural	characteristics	it	produces;	sociology	investigates	the	human kind	in	terms	of	social	relations	and	institutions,	cultural	anthropology	is	the	comparative	study	of	these matters,	and	so	forth. 3.	A	third	desideratum	concerns	the	relationship	between	a	new	conception	of	human	nature	and	the existing,	vernacular	conception.	The	new	conception	cannot	include	as	defining	conditions	of	human nature	all	the	features	that	are	associated	with	the	vernacular	concept.	As	we	have	already	mentioned, these	are	essentially	independent	biological	properties	that	we	should	not	expect	to	be	tightly	associated with	each	other	(Griffiths	2011).	But	there	are	important	properties	that	some	human	phenotypes exhibit,	and	a	concept	of	human	nature	should	recognise	this.	For	example,	the	fixity	of	traits	can	be explained	by	canalisation	(Waddington	1942)	and	the	fact	that	there	are	canalised	traits	should	be	part of	our	understanding	of	human	nature.	Typicality	is	not	a	defining	feature	of	human	nature,	but	the	fact that	there	are	some	typical	features	of	human	beings	needs	to	be	encompassed	by	our	understanding	of human	nature.	Teleology	is	today	standardly	explained	via	evolutionary	adaptation	–	some	features really	are	there	by	evolutionary	'design'	and	others	are	not	–	so	our	understanding	of	human	nature should	recognise	that	our	nature	is	in	part	the	outcome	of	evolutionary	design3. 4.	Finally,	contra	Edouard	Machery	(Machery	2008),	universality	is	not	a	desideratum	for	a	concept	of human	nature	(see	Ramsey	2012;	Ramsey	2013	for	a	critique).	If	the	human	species	is	polymorphic,	then this	is	part	of	the	nature	of	the	human	species,	something	we	should	seek	to	understand	when	we	study human	beings	as	a	kind.	Many	organisms	also	exhibit	some	form	of	phenotypic	plasticity,	the	evolved ability	to	respond	with	different	phenotypes	to	different	environments	(Gilbert	and	Epel	2009;	Sultan 2015).	This	too	is	an	important	part	of	the	nature	of	the	species	in	question. In	suggesting	that	the features	of	human	nature	must	be	universal	Machery	is	responding	to	a	real	feature	of	the	vernacular concept	of	human	nature,	but	one	that	clashes	with	what	we	have	learnt	about	biology	since	Darwin.	So our	fourth	desideratum	is	that	human	nature	should	admit	of	polymorphism	and	plasticity. In	summary,	then,	we	propose	that	a	concept	of	human	nature	should	make	human	nature	something that	explains	many	features	of	human	beings,	that	it	should	make	human	nature	an	object	of	enquiry	for the	human	sciences,	that	human	nature	should	make	room	for	developmentally	fixed	and	species-typical traits,	and	for	the	fact	that	some	traits	are	the	result	of	evolutionary	design,	and	finally	that	it	is	part	of human	nature	that	humans	are	plastic	and	variable. Amongst	the	many	accounts	of	human	nature	offered	by	philosophers	two	meet	these	desiderata.	The first	is	Grant	Ramsey's	Life-History	Trait	Cluster	(LTC)	account	of	human	nature	(Ramsey	2013)	and	the second	is	the	Developmental	Systems	(DS)	account	of	human	nature	(Stotz	2010;	Griffiths	2011).	In	the next	section	we	explain	the	similarities	and	complementarities	of	these	two	accounts,	and	in	Section	Five we	turn	to	the	differences	between	them. 3	One	problematic	aspects	of	the	teleological	way	of	thinking	is	its	resistance	to	counter	evidence.	The	nonexistence	of	a	so-called	essential	trait	among	a	large	number	of	members	of	a	population	can	always	be	explained as	the	failure	of	those	individuals	to	realise	their	proper	nature	(we	thank	to	the	editors	for	this	comment). 4. LTC	and	DST:	Human	nature	as	human	development So	what	is	the	LTC	account?	Ramsey	acknowledges	that	human	beings	are	diverse,	with	each	individual life	history	including	a	different	mix	of	traits.	His	account	focuses	on	the	patterns	of	co-occurrence between	traits	in	this	population	of	diverse	life	histories	(Ramsey	2013,	987). Human	nature	is	defined	as	the	pattern	of	trait	clusters	within	the	totality	of	extant	human possible	life	histories.	Thus,	if	one	were	to	take	all	of	the	possible	life	histories	that	form	the	basis for	individual	nature,	and	then	combine	them,	one	would	possess	the	set	of	life	histories	that forms	the	basis	for	human	nature,	since	the	trait	distribution	patterns	in	this	set	of	life	histories constitute	human	nature.	(Ramsey	2013,	987) Two	ideas	are	combined	in	this	proposal	both	of	which	are	central	to	developmental	systems	theory: first,	"from	an	evolutionary	point	of	view	an	animal	is	the	implementation	of	a	life-history	strategy"; second,	"bringing	order	to	that	diversity	is	not	about	identifying	universal	elements,	but	about	finding order	in	the	patterns	of	similarity	and	difference"	(Griffiths	2011,	325	&	328).	In	fact,	as	we	now	go	on	to show,	the	two	accounts	are	remarkably	convergent,	albeit	arriving	at	their	conclusions	from	very different	starting	points. Ramsey	identifies	the	key	desiderata	for	an	account	of	human	nature	account	to	both	with	scientific practice	and	with	intuitive	notions	of	human	nature.	The	first	demands	its	empirical	accessibility	as	a subject	to	the	human	sciences,	which	is	in	line	with	our	second	desideratum.	Ramsey	also	wants	his account	of	human	nature	to	clarify	the	related	concepts	of	"innateness	and	naturalness"	(Ramsey	2013, 986).	This	requirement	has	something	in	common	with	our	third	desiderata,	that	a	concept	of	human nature	should	shed	light	on	the	phenomena	of	typicality,	fixity	and	teleology.	Ramsey's	account	also embraces	developmental	plasticity	and	diversity,	and	so	meets	our	fourth	desideratum. There	are	other	similarities	between	the	two	accounts.	Ramsey	eschews	any	classificatory	role	for human	nature:	an	organism	is	human	because	it	is	a	member	of	a	particular	lineage,	not	because	it displays	the	LTC	property	clusters.	He	also	eschews	a	normative	role	for	human	nature:	his	account	may illuminate	the	idea	that	some	traits	are	'natural',	but	it	is	not	intended	as	an	account	of	how	human beings	should	be.	We	agree	with	both	of	these	points. Ramsey	seems	to	regard	his	account	as	merely	descriptive	and	not	explanatory:	"characterizations	of features	of	human	nature	are	merely	descriptions	of	patterns	within	the	collective	set	of	human	life histories"	(Ramsey	2013,	988).	This	apparently	clashes	with	our	first	desideratum,	which	calls	for	an explanatory	account	of	human	nature. For	two	reasons,	however,	we	do	not	see	this	as	a	major difference	between	our	accounts.	First,	Ramsey's	life-history	trait	clusters	are	exactly	what	our developmental	systems	account	of	human	nature	explains,	which	makes	his	account	complementary	to ours.	Secondly,	we	believe	that	Ramsey	actually	presents	an	account	of	how	individual	traits	that	make up	the	trait	cluster	can	be	explained	by	human	nature. Ramsey's	account	is	more	than	merely	descriptive,	we	think,	because	it	does	not	simply	list	features	as	a description	of	human	nature.	The	account	focuses	on	the	identification	of	'antecedent'	(A)	and 'consequent'	(C)	traits	of	life	histories	that	have	been	found	to	be	associated	with	each	other.	Further experiments	should	then	be	carried	out,	Ramsey	suggests,	to	determine	if	As	and	Cs	are	causally	related rather	than	merely	correlated.	This	would	amount	to	an	experimental	programme	to	establish constraints	on	the	possible	trajectories	within	life-history	space,	and	hence	the	beginning	of	an explanation	of	human	nature	as	well	as	a	description. Ramsey	sketches	a	quasi-formal	account	of	human	nature,	involving	a	'human	nature	space'.	This	is	not the	state	space	of	human	life	histories,	as	in	the	last	paragraph,	but	a	theoretical	space	in	which	to	locate and	compare	particular	traits	clusters.	It	has	two	dimensions:	the	"pervasiveness,	p,	of	the	antecedent", defined	as	"the	proportion	of	life	histories	that	exhibit	that	trait",	and	the	"robustness,	r,	of	the antecedent-consequent	association"	(we	are	unclear	if	r	is	simply	a	correlation,	something	like	the regression	of	the	consequent	on	the	antecedent,	or	more	explicitly	a	causal	measure).	One	can	increase	p by	choosing	a	more	broadly	defined	antecedent	but	this	will	typically	reduce	the	robustness	of	its association	with	a	consequent.	Equally,	adding	more	antecedent	traits	make	it	more	complex	can increase	r,	but	at	a	cost	to	p.	Hence	there	is	a	trade-off	between	p	and	r,	or	between	simplicity	and strength	(Ramsey	2013,	989-990). Ramsey	argues	that	one	can	make	sense	of	both	innateness	and	naturalness	in	terms	of	positions	within the	p-r	space.	It	may	be	natural,	part	of	human	nature,	for	humans	that	have	property	A	to	also	have property	C.,	for	example,	being	female	(A)	and	menstruating	(C).	Since	human	nature	is	also	associated with	traits	being	innate,	innateness	could	be	interpreted	in	various	ways	in	terms	of	the	p-r	space.	Either the	higher	the	r-value,	the	more	innate	a	trait	is;	or	innateness	can	be	defined	as	associations	with	both	a high	p-value	and	a	high	r-value;	or,	since	neither	of	these	two	proposed	definitions	of	innateness	imply 'not	learned'	one	could	restrict	the	term	innate	to	A-C	links	that	don't	involve	any	learning.	This,	Ramsey admits,	might	exclude	most,	if	not	all,	associations	"since	learning	is	woven	into	the	causal	fabric	of	so much	of	development"	(Ramsey	2013,	991). Ramsey	notes	that	an	LTC	account	of	human	nature	may	seem	"spectacularly	–	and	perhaps	disastrously –	permissive"	and	"extremely	inclusive"	(Ramsey	2013,	987).	However,	Ramsey	argues	that	although	LTC is	in	principle	very	permissive	in	that	it	includes	all	trait	associations,	it	does	not	imply	that	all	these associations	are	equally	interesting.	He	proposes	the	p-r	space	as	a	way	of	to	distinguish	the	more interesting	features	of	human	nature,	those	most	worthy	of	study	in	science	of	human	nature.	In	so	far	as these	interesting	trait	associations	are	the	ones	that	are	more	'natural'	or	'innate',	this	seems	to	us	to	be another	residual	influence	of	the	folkbiological	conception	of	human	nature.	In	any	case,	Ramsey	does not	need	to	defend	himself	against	this	criticism.	As	we	explain	in	the	next	section,	the	idea	that	accounts of	human	nature	should	not	be	'permissive'	or	'inclusive'	is	simply	mistaken. Finally,	Ramsey	claims	that	while	his	account	is	not	normative,	it	nevertheless	has	"moral	implications". Since	this	account	gives	us	robust	insight	into	the	human	condition,	good	and	bad,	it	could	guide	action via	desired	or	unwanted	antecedent-consequent	associations	(Ramsey	2013,	992).	That	biology	can	have moral	implications	in	this	straightforward	way	has	often	been	noticed:	"Starving	children	stunts	their growth	and	ruins	their	health	and	this	is	one	reason	not	to	starve	them"	(Sterelny	and	Griffiths	1999,	5). The	Developmental	Systems	Account	of	Human	Nature The	developmental	systems	account	describes	human	nature	in	a	way	very	similar	to	Ramsey.	Organisms are	fundamentally	processes	(Griffiths	and	Stotz	In	Press)	life	cycles	and	heredity	is	the	reconstruction of	the	life	cycle	using	resources	that	are	passed	on	by	previous	generations.	Some	of	these	resources	are genetic,	some	epigenetic.	and	some	exogenetic,	the	last	term	referring	to	a	'developmental	niche'	that contains	reliable	developmental	resources	from	outside	the	organisms.4	Some	exogenetic	resources serve	to	canalise	development,	and	some	to	modify	it	and	hence	enable	developmental	plasticity. Developmental	systems	theorists	have	long	recognised	Ramsey's	point	that	a	single	lineage	has	many possible	developmental	trajectories:	"life	cycles	may	have	a	disjunctive	form,	with	different	individuals having	different	characteristics.	A	developmental	system	can	proliferate	by	producing	a	range	of outcomes	on	different	occasions"	(Griffiths	and	Gray	1994,	296).	Descriptive	human	nature	is	the	"order in	the	patterns	of	similarity	and	difference"	in	these	human	life	cycles	(Griffiths	2011,	328). Developmental	systems	theory	explains	human	nature	as	the	product	of	the	human	developmental system,	a	matrix	of	genetic,	epigenetic,	and	exogenetic	resources	within	which	the	developmental process	or	life	cycle	unfolds.	This	system	is	constructed	by	earlier	human	life	cycles	and	by	feed-forward 4	For	an	account	of	the	unique	features	of	human	nature	that	accords	a	very	substantial	role	to	the developmental	niche,	see	Sterelny	2013;	Sterelny	2012. effects	from	the	development	of	the	individual	itself.	Progress	in	understanding	human	nature,	on	this view,	is	simply	progress	in	the	sciences	of	human	development:	developmental	biology,	developmental psychobiology,	and	developmental	psychology	(for	a	brief	history	of	DST	and	the	scientific	research traditions	from	which	it	emerged,	see	Griffiths	and	Tabery	2013). Ramsey's	fear	that	the	LTC	account	might	be	criticised	as	"spectacularly	–	and	perhaps	disastrously	– permissive"	echoes	Kronfeldner	and	collaborators	assessment	of	the	developmental	systems	account	of human	nature: The	result	is	a	concept	of	an	all-inclusive	human	nature	that	comprises	all	the	resources	needed	to stabilize	the	development	of	the	patterns	of	similarity	and	difference	observable	in	humankind. Human	nature,	the	thing	that	explains	and	defines	the	human	species,	is	then	a	genealogically anchored	explanatory	essence	of	gigantic	proportions,	namely	the	whole	developmental	system	of humankind,	including	the	developmental	niche	... [this	is]	a	very	distant	relative	of	the	traditional	concept	of	human	nature,	since	it	construes everything	involved	in	and	resulting	from	human	development	as	part	of	human	nature.	It	is doubtful	whether	such	an	all-encompassing	concept	of	human	nature	is	of	any	concrete	use	for the	sciences,	that	is,	for	describing	and	explaining	commonalities	or	explaining	differences	within humankind	or	between	the	human	and	other	species	(Kronfeldner,	Roughley,	and	Toepfer	2014, 649,	our	italics). This	is	a	non-sequitur:	the	observations	made	about	the	DST	account	do	not	support	the	conclusion.	In fact,	the	DST	account	has	a	better	prospect	of	"describing	and	explaining	commonalities	or	explaining differences"	than	the	"traditional	concept	of	human	nature"	which	excludes	much	of	human	diversity.	If the	aim	was	to	pick	out	some	individuals	as	not	human	or	as	less	human,	then	we	might	need	a	simple definition	of	the	human,	like	a	CO1	gene	barcode,	but	hopefully	no-one	is	trying	to	do	that! If,	instead,	the aim	of	studying	human	nature	is	to	understand	what	human	beings	are	like	and	why	they	are	like	that, then	we	see	no	reason	why	either	the	description	of	human	nature	or	its	underlying	explanation	should be	simple.	It	seems	obvious	that	both	will	be	complex. Kronfeldner	et	al.	seem	to	be	echoing	a	common	criticism	levelled	at	developmental	systems	theory, namely	that	paying	attention	to	the	role	of	the	environment	in	development	and	to	the	plasticity	of development	will	make	the	study	of	development	scientifically	intractable	and	its	results incomprehensibly	complex.	The	same	accusation	has	been	levelled	against	the	scientists	whose	work inspired	DST,	and	the	reply	is	the	one	those	scientists	gave	–	"development	is	complicated"	(Bateson 1991,	19,	italics	in	original).	Complex	interactions	between	genetic,	epigenetic	and	exogenetic	factors explain	the	constraints	on	developmental	trajectories	in	the	state	space	of	possible	human	life	histories that	constitute	human	nature	on	our	view,	and	on	Ramsey's	view	as	we	interpret	it5.	It	would	be convenient	if	these	could	be	reduced	to	a	few	simple	parameters,	like	the	average	velocity	of	molecules	in a	gas,	but	it	is	clear	that	they	cannot.6 Developmental	systems	theorists	have	repeatedly	emphasised	that	an	inclusive	definition	of	the developmental	system	does	not	mean	that	the	whole	system	must	be	studied	at	once,	any	more	than	the inclusive	definition	of	the	proteome	precludes	studying	individual	protein-protein	interactions	(Griffiths and	Gray	2005;	Oyama	2000).	The	concept	of	the	developmental	niche,	which	seems	to	be	of	particular concern	to	Kronfeldner	et	al.,	is	a	construct	from	empirical	research	on	behavioural	development	(West and	King	1987)	that	was	introduced	into	DST	to	give	greater	structure	to	the	extra-organismic component	of	the	developmental	system	(Stotz	and	Allen	2012;	Stotz	and	Griffiths	2016).	The developmental	niche	concept	has	been	used	to	great	effect	in	such	different	fields	as	the	development	of social	behaviour	and	communication	in	birds	(West	and	King	1987;	West	and	King	2008),	and	speciestypical	development	in	general	in	rats	(Alberts	2008).	Other	research	groups	have	applied	DST's	view	of development	and	the	concept	of	the	developmental	niche	to	investigate	aspects	of	human	development (Alberts	and	Ronca	2012;	Gros-Louis,	West,	and	King	2014;	Gros-Louis,	West,	and	King	2016;	Narvaez	et al.	2013).	None	of	this	research	has	become	mired	in	an	unmanageable	sea	of	complexity	because	it recognises	that	the	human	life-cycle	has	evolved	to	make	use	of	a	highly	specific	developmental	niche,	or that	interaction	with	this	niche	may	induce	developmental	plasticity. Developmental	systems	theory	does	not	make	it	possible	to	sum	up	human	nature	in	a	slogan,	but	it	does point	clearly	to	the	body	of	knowledge	that	constitutes	our	current	best	understanding	of	human	nature: human	developmental	biology,	developmental	psychobiology	and	developmental	psychology.	When those	sciences	are	complete	we	will	have	a	complete	understanding	of	human	nature.	We	fail	to	see	the force	of	the	objection	'but	that	will	be	very	complicated'. 5	A	difference	between	Ramsey's	account	and	our	own	may	concern	the	status	of	the	genome,	which	DST put	on	a	much	more	equal	footing	to	the	environment	than	it	enjoys	on	the	LTC	account:	"If	genes	were allowed	to	vary,	individual	nature	would	be	vacuous	since	sufficient	changes	to	genes	could,	say,	change an	American	into	an	aardvark.	By	contrast,	varying	the	way	that	an	individual	encounters	its environmental	heterogeneity	reveals	something	about	its	nature"	(Ramsey	this	volume). 6	This	is	not	to	reject	research	programs	in	systems	biology	that	aim	at	substantial	reductions	in	the complexity	of	development	through	identifying	systems-level	variables. 5.	A	distinctive	feature	of	the	DS	account	human	developmental	niche	construction While	Ramsey	focuses	on	descriptive	property	clusters	that	make	up	human	nature,	the	developmental systems	account	focuses	on	the	underlying	processes	that	account	for	these	clusters.	Developmental systems	theory	subscribes	to	a	process	account	of	the	organism,	and	this	is	reflected	in	its	view	of	human nature.	DST	is	a	process	theory	because	developmental	systems	are	essentially	extended	in	time (Griffiths	and	Gray	1994;	Griffiths	and	Gray	1997;	Griffiths	and	Stotz	In	Press).	DST	"seeks	to	explain developmental	outcomes	as	the	result	of	a	dynamic	process	in	which	some	of	the	interacting	factors	are products	of	earlier	stages	of	the	process,	rather	than	as	the	result	of	the	arrangement	of	pre-existing factors	into	a	static	mechanism.	Even	when	factors	exist	independently	of	the	developmental	process, they	are	drawn	into	it	and	made	part	of	a	developmental	'system'	by	the	unfolding	process"	(Griffiths	and Stotz	In	Press).	The	focus	on	property	clusters	makes	Ramsey's	LTC	account	look	less	processual. However,	these	properties	are	not	merely	properties	of	organisms,	but	properties	of	an	organism	at	a time,	and	the	property	clusters	that	constitute	human	nature	are	correlations	between	what	happens	at one	point	in	a	life	cycle	and	what	happens	at	a	later	point.	So	Ramsey's	account	actually	fits	a	process view	of	the	organism	quite	well.	Moreover,	Ramsey	conceives	of	the	series	of	events	that	make	up	an individual	human	being	as	a	life-history,	the	implementation	of	an	evolved	strategy	for	resource allocation	across	the	lifespan.	In	our	recent	work	we	have	argued	that	it	is	a	life-history	strategy	that constitutes	the	principle	of	identity	that	makes	a	series	of	events	into	a	single	life-cycle,	rather	than	a	part of	a	larger	cycle,	or	a	process	involving	more	than	one	individual	(Griffiths	and	Stotz	In	Press). The	main	difference	between	the	two	accounts	is	that	the	DS	account	has	a	stronger	focus	on	the	role	of the	environment	in	constituting	human	nature.	There	is	an	old	saying	within	anthropology	that	culture	is not	only	part	of	human	nature,	but	that	our	nature	is	culture.	Some	recent	work	on	human	evolution	has emphasised	the	role	of	selective	niche	construction:	the	evolution	of	the	unique	characteristics	of	human psychology	and	social	structure	has	been	substantially	driven	by	the	selection	pressures	created	by earlier	psychologies	and	social	structures	(Laland,	Odling-Smee,	and	Feldman	2000;	Sterelny	2012). Niche	construction	theory	deals	with	the	selective	niche,	defined	by	the	parameters	that	determine	the relative	fitness	of	competing	types	in	a	population.	In	selective	niche	construction	earlier	generations partly	construct	the	selection	pressures	that	act	on	future	generations. But	another	aspect	of	human niche	construction	is	that	our	development	is	dependent	on	a	rich	developmental	niche	of	interaction with	parents	and	other	conspecifics	and	with	physical	and	cognitive	artifacts	from	tools	to	languages. The	developmental	niche	is	defined	by	the	parameters	needed	to	ensure	the	reconstruction	of	the evolved	life	cycle.	The	concept	of	the	developmental	niche	is	designed	to	integrate	and	formalize	the	nongenetic	yet	heritable	factors	influencing	an	organism's	development	(Stotz	2010;	Stotz	2014;	Stotz	Under Review;	Griffiths	and	Stotz	2013).	The	concept	goes	back	to	developmental	psychobiologists	West	and King	(West	and	King	1987). In	our	current	formulation	of	the	concept	(Griffiths	and	Stotz	2013;	Griffiths	and	Stotz	In	Press),	the developmental	system	consists	of	genetic	resources,	epigenetic	resources	and	an	exogenetic developmental	niche,	which	contains	reliably	inherited	physical,	social,	ecological	and	epistemic resources	needed	to	reconstruct,	or	in	the	case	of	phenotypic	plasticity	to	modify,	that	developmental system.	These	resources	can	be	actively	constructed	by	the	parents	(producing	the	'parental	effects'	of quantitative	genetics)	or	by	the	larger	group,	co-constructed	by	parent	and	offspring,	or	sourced passively	from	a	stable	environment.	Wherever	they	come	from,	if	there	exists	an	evolutionary (historical)	explanation	for	the	interaction	of	the	evolved	developmental	system	with	the	resource	then that	resource	is	part	of	the	system.	What	evolves	by	natural	selection	is	a	relationship	between	system and	each	resource. How	does	the	developmental	niche	influence	human	development?	Human	babies	are	needy.	They	are born	early	in	comparison	to	other	primates,	meaning	that	for	several	months	postnatally,	relative	to other	primates,	human	babies	share	characteristics	of	fetuses	rather	than	of	infants	in	those	other primates	(Trevathan	2011).	Comparing	brain	size	at	birth	among	primates,	humans	should	be	born	at	18 months	of	age.	A	large	part	of	brain	development	takes	place	outside	the	uterus,	allowing	for	much greater	postnatal	epiand	exo-genetic	influence	than	for	their	ape	cousins,	which	makes	the	early	niche fundamental	for	human	development.	Over	the	course	of	human	evolution,	as	brains	became	bigger	and human	infants	more	immature	at	birth,	human	childrearing	practices	evolved	in	tandem	with	these changes	to	ensure	the	survival	of	the	helpless	infant.	As	bipedalism,	hemochorial	placenta,	large	brains and	the	need	for	a	greater	amount	of	learning	after	birth	emerged,	human	evolution	intensified	parental care:	"Only	with	intensified	parental	care	in	response	to	greater	helplessness	of	the	infant	could	selection favor	the	evolution	of	a	large	brain	in	a	bipedal	animal"	(Trevathan	2011,	33).	The	evolution	of	a	more complex	and	resource-demanding	developmental	niche	has	been	a	key	feature	of	human	evolution. For	this	reason,	it	seems	to	us	entirely	natural	to	say	that	that	human	nature	resides	partly	in	the	human developmental	environment.	We	are	a	species	that	is	particularly	strongly	influenced	by	niche construction,	both	selective	niche	construction	over	evolutionary	timescales	and	developmental	niche construction	over	ontogenetic	timescales. A	concept	of	nature	according	to	which	what	is	natural	must come	from	the	inside	is	particularly	unsuitable	for	such	a	species.	Imagine	trying	to	determine	the	real nature	of	an	ant,	another	powerful	niche	constructor,	by	removing	the	influence	of	the	nest	on	the developing	egg	and	embryo.	The	result	would	be	either	dead	or	biologically	meaningless,	and	so	it	is	for humans. The	developmental	niche	has	two	fundamental	functions.	One	function	is	to	ensure	the	stable,	reliable development	of	species-typical	traits.	So	what	explains	Typicality	is	the	developmental	systems	dynamics within	what	we	may	call	'normal'	parameters,	some	of	which	are	provided	by	pre-existing	physical	and developmental	constraints.	The	rest	are	ensured	by	reliably	and	stably	inherited	resources,	which include	not	just	the	genome	but	also	essential	environmental	resources	that	assist,	among	other functions,	in	the	species-typical	expression	of	the	genetic	factors.	These	stable	resources	also	partially explain	fixity.	In	addition	there	are	developmental	mechanisms	that	buffer	against	internal	(genetic, epigenetic,	metabolic)	and	external	perturbations.	These	are	invoked	when	we	talk	about	canalization. But	one	of	our	desiderata	was	that	human	nature	needs	to	embrace	and	explain	human	diversity:	"The search	for	a	shared	human	nature	cannot	be	the	search	for	human	universals;	it	must	instead	be	a	way	to interpret	and	make	sense	of	human	diversity"	(Griffiths	2011,	326).	Here	the	second	function	of	the developmental	niche	comes	in.	Beyond	ensuring	reliable	development,	the	developmental	niche	also provides	input	to	developmental	plasticity.	Plasticity	is	often	defined	in	terms	of	a	genotype's	ability	to produce	different	phenotypes	in	response	to	the	environment.	It	would	be	more	accurate,	however,	to say	that	the	shape	of	the	norm	of	reaction	is	a	property	of	the	whole	developmental	system.	So	what explains	human	diversity	are	differing	developmental	systems	dynamics	supported	by	modifications	in the	developmental	niche.	In	other	words,	human	diversity	results	primarily	from	the	interaction	between the	evolved	developmental	system	and	a	wide	range	of	environments,	including	novel	environments. "Bringing	order	to	that	diversity	is	not	about	identifying	universal	elements,	but	about	finding	order	in the	patterns	of	similarity	and	difference"	(Griffiths	2011,	328).	Developmental	niche	construction therefore	provides	dependability,	but	also	adaptive	flexibility,	in	the	provision	of	necessary developmental	resources. 6.	Conclusion In	this	chapter	we	have	reiterated	our	view	that	human	nature	is	simply	human	development.	To	the extent	that	we	understand	human	developmental	biology,	developmental	psychobiology	and developmental	psychology,	we	understand	human	nature.	Like	Ramsey's	LTC	account	of	human	nature, this	amounts	to	saying	that	human	nature	is	a	set	of	constraints	on	possible	human	developmental trajectories.	Like	Ramsey's	account,	it	is	not	without	content	because,	although	it	does	not	identify	a	set of	outcomes	that	are	'unnatural',	it	does	say	that	'you	can't	get	there	from	here'.	This	gives	our	account, like	Ramsey's,	a	very	special	and	positive	feature,	namely	that	it	is	able	to	embrace	human	diversity	as part	of	human	nature.	As	we	have	argued	in	section	two,	the	objection	that	our	account	leads	to	a	very complex	picture	of	human	nature	is	a	non	sequitur:	human	nature	is	complicated. Our	account	differs	from	Ramsey's	in	a	greater	stress	on	the	role	of	the	human	developmental environment	–	the	developmental	niche	–	in	constituting	human	nature.	We	have	argued	that	this	reflects the	direction	of	the	human	sciences	in	recent	years.	If	it	clashes	with	a	folkbiological	intuition	that	nature must	come	from	'inside;'	rather	than	'outside',	so	much	the	worse	for	that	intuition	–	we	understand human	nature	better	now. Kronfeldner	et	al	(Kronfeldner,	Roughley,	and	Toepfer	2014;	Kronfeldner	In	Press)	have	proposed	a 'pluralistic	solution'	for	the	missing	consensus	in	the	philosophical	literature	regarding	a	concept	of human	nature.	They	suggest	that,	"different	scientific	fields	are	in	need	of	different	concepts	of	human nature,	each	fulfilling	an	independent	epistemic	role."	We	are	sympathetic	to	this	general	approach	to	the analysis	of	scientific	concepts.	We	ourselves	have	made	a	similar	suggestion	about	the	concept	of	the gene:	different	gene	concepts	should	be	understood	as	"tools	of	research,	as	ways	of	classifying	the experience	shaped	by	experimentalists	to	meet	their	specific	needs"	(Stotz	and	Griffiths	2008;	see	also Griffiths	and	Stotz	2013).	We	do	not,	however,	think	that	the	same	argument	applies	to	the	concept	of human	nature.	Human	nature	is	much	less	a	technical	concept	applied	in	the	laboratory	than	a	pragmatic, and	even	normative,	tool	applied	in	wider	social	contexts	and	with	wide-ranging	consequences.	This does	not	mean	that	different	scientific	endeavours	cannot	study	different	aspect	of	human	nature,	but they	cannot	do	this	without	paying	attention	to	other	fields.	There	have	been	several	attempts	to	impose a	simplistic	understanding	of	human	nature,	often	derived	from	evolutionary	biology,	and	to	marginalise other	sciences,	such	as	those	that	focus	on	the	human	developmental	environment.	As	Sandra	Mitchell has	argued,	an	'anything	goes'	pluralism	in	science	may	do	more	harm	than	good,	while	a	real	'integrative pluralism'	is	a	useful	alternative	against	a	reductionist	imperialism	(Mitchell	2003;	Mitchell	2009). References Alberts,	Jeffrey	R.	2008.	"The	Nature	of	Nurturant	Niches	in	Ontogeny."	Philosophical	Psychology	21	(3): 295–303. Alberts,	Jeffrey	R.,	and	April	E.	Ronca.	2012.	"The	Experience	of	Being	Born:	A	Natural	Context	for Learning	to	Suckle."	International	Journal	of	Pediatrics	2012	(September):	e129328. doi:10.1155/2012/129328. Atran,	Scott.	1990.	Cognitive	Foundations	of	Natural	History:	Towards	an	Anthropology	of	Science. Cambridge	and	New	York:	Cambridge	University	Press. Bateson,	Patrick	P.	G.	1991.	"Are	There	Principles	of	Behavioural	Development?"	In	The	Development	and Integration	of	Behaviour:	Essays	in	Honour	of	Robert	Hinde,	edited	by	Patrick	P	G	Bateson,	19–39. Cambridge:	Cambridge	University	Press. Berlin,	Brent.	1992.	Ethnobiological	Classification:	Principles	of	Classification	of	Plants	and	Animals	in Traditional	Societies.	Princeton,	NJ:	Princeton	University	Press. Cheung,	Benjamin	Y.,	Ilan	Dar-Nimrod,	and	Karen	Gonsalkorale.	2014.	"Am	I	My	Genes?	Perceived Genetic	Etiology,	Intrapersonal	Processes,	and	Health:	Am	I	My	Genes?"	Social	and	Personality Psychology	Compass	8	(11):	626–37. Dar-Nimrod,	Ilan,	and	Steven	J.	Heine.	2011a.	"Genetic	Essentialism:	On	the	Deceptive	Determinism	of DNA."	Psychological	Bulletin	137	(5):	800–818. ---.	2011b.	"Some	Thoughts	on	Essence	Placeholders,	Interactionism,	and	Heritability:	Reply	to Haslam	(2011)	and	Turkheimer	(2011)."	Psychological	Bulletin	137	(5):	829–33. Dar-Nimrod,	Ilan,	and	Grace	Lisandrelli.	2012.	"It's	in	My	Genes.	Causal	Attributions	and	Perceptions	of Choice."	Advances	in	Psychology	Research	95:	187–98. Dar-Nimrod,	Ilan,	Miron	Zuckerman,	and	Paul	Duberstein.	2014.	"Smoking	at	the	Workplace:	Effects	of Genetic	and	Environmental	Causal	Accounts	on	Attitudes	toward	Smoking	Employees	and Restrictive	Policies."	New	Genetics	and	Society	33	(4):	400–412. Dar-Nimrod,	Ilan,	Miron	Zuckerman,	and	Paul	R.	Duberstein.	2012.	"The	Effects	of	Learning	about	One's Own	Genetic	Susceptibility	to	Alcoholism:	A	Randomized	Experiment."	Genetics	in	Medicine	15: 132–38. Downes,	Stephen	M.,	and	Edouard	Machery.	2013.	"Arguing	about	Human	Nature:	Contemporary Debates."	http://philpapers.org/rec/DOWAAH. Fuentes,	A.,	J.	Marks,	T.	Ingold,	and	R.	Sussman.	2010.	"On	Nature	and	the	Human."	American Anthropologist	112	(4):	512–21. Gelman,	S.	A.	(2003).	The	essential	child:	Origins	of	essentialism	in	everyday	thought.	New	York:	Oxford University	Press. Gilbert,	Scott,	and	David	Epel.	2009.	Ecological	Developmental	Biology:	Integrating	Epigenetics,	Medicine, and	Evolution.	Sunderland,	MA:	Sinauer	Associates. Glackin,	Shane	N.	2015.	"Three	Aristotelian	Accounts	of	Disease	and	Disability."	Journal	of	Applied Philosophy.	doi:10.1111/japp.12114. Gottlieb,	Gilbert.	1970.	"Conceptions	of	Prenatal	Behavior."	In	Development	and	Evolution	of	Behavior: Essays	in	Memory	of	T.	C.	Schneirla,	edited	by	L.	R	Aronson,	E	Tobach,	D.	S	Lehrman,	and	J.	S Rosenblatt,	111–37.	San	Francisco:	W.	H.	Freeman. Griffiths,	Paul	E.	2011.	"Our	Plastic	Nature."	In	Transformations	of	Lamarckism: From	Subtle	Fluids	to Molecular	Biology,	edited	by	Snait	Gissis	and	Eva	Jablonka,	319–30.	Cambridge,	MA:	MIT	Press. Griffiths,	Paul	E.	2002.	"What	Is	Innateness?"	The	Monist	85	(1):	70–85. Griffiths,	Paul	E.,	and	Russell	D.	Gray.	1994.	"Developmental	Systems	and	Evolutionary	Explanation." Journal	of	Philosophy	XCI	(6):	277–304. ---.	1997.	"Replicator	II:	Judgment	Day."	Biology	and	Philosophy	12	(4):	471–92. ---.	2005.	"Three	Ways	to	Misunderstand	Developmental	Systems	Theory."	Biology	&	Philosophy	20 (2):	417–25. Griffiths,	Paul	E.,	and	Edouard	Machery.	2008.	"Innateness,	Canalisation	and	'Biologicizing	the	Mind.'" Philosophical	Psychology	21	(3):	397–414. Griffiths,	Paul	E.,	Edouard	Machery,	and	Stefan	Linquist.	2009.	"The	Vernacular	Concept	of	Innateness." Mind	and	Language	24	(5):	605–30. Griffiths,	Paul	E.,	and	Karola	Stotz.	2013.	Genetics	and	Philosophy:	An	Introduction.	New	York:	Cambridge University	Press. Griffiths,	Paul	E.,	and	Karola	Stotz.	In	Press.	"How	DST	Became	a	Process	Theory."	In	Everything	Flows: Towards	a	Processual	Philosophy	of	Biology,	by	Daniel	J.	Nicholson	and	John	Dupre.	Oxford	and New	York:	Oxford	University	Press. Griffiths,	Paul	E.,	and	James	G.	Tabery.	2013.	"Developmental	Systems	Theory:	What	Does	It	Explain,	and How	Does	It	Explain	It?"	In	Embodiment	and	Epigenesis:	Theoretical	and	Methodological	Issues	in Understanding	the	Role	of	Biology	within	the	Relational	Developmental	System	Part	A:	Philosophical, Theoretical,	and	Biological	Dimensions,	edited	by	Richard	M	Lerner	and	Janette	B	Benson,	45:65– 94.	Advances	in	Child	Development	and	Behavior	45.	Elsevier. Gros-Louis,	Julie,	Meredith	J.	West,	and	Andrew	P.	King.	2014.	"Maternal	Responsiveness	and	the Development	of	Directed	Vocalizing	in	Social	Interactions."	Infancy	19	(4):	385–408. ---.	2016.	"The	Influence	of	Interactive	Context	on	Prelinguistic	Vocalizations	and	Maternal Responses."	Language	Learning	and	Development	12	(3):	280. Hinde,	R.	A.	1968.	"Dichotomies	in	the	Study	of	Development."	In	Genetic	and	Environmental	Influences	on Behaviour,	edited	by	J.M	Thoday	and	A.S	Parkes,	3–14.	New	York:	Plenum. Hull,	D.	L.	1986.	"On	Human	Nature."	Proceedings	of	the	Philosophy	of	Science	Association	Vol.1:	3–13. Jayaratne,	Toby	Epstein,	Susan	A	Gelman,	Merle	Feldbaum,	Jane	P	Sheldon,	Elizabeth	M	Petty,	and	Sharon L.	R	Kardia.	2009.	"The	Perennial	Debate:	Nature,	Nurture,	or	Choice?	Black	and	White	Americans' Explanations	for	Individual	Differences."	Review	of	General	Psychology	13	(1):	24–33. Kronfeldner,	Maria.	In	Press.	"Divide	and	Conquer:	The	Authority	of	Nature	and	Why	We	Disagree	about Human	Nature."	In	Why	We	Disagree	about	Human	Nature,	edited	by	Tim	Lewens	and	Beth Hannon.	Oxford	University	Press. Kronfeldner,	Maria,	Neil	Roughley,	and	Georg	Toepfer.	2014.	"Recent	Work	on	Human	Nature:	Beyond Traditional	Essences."	Philosophy	Compass	9	(9):	642–52. Laland,	K.	N,	F.	John	Odling-Smee,	and	M.W	Feldman.	2000.	"Niche	Construction,	Biological	Evolution	and Cultural	Change."	Behavioral	and	Brain	Sciences	23:	131–57. Lewens,	Tim.	2012.	"Human	Nature:	The	Very	Idea."	Philosophy	and	Technology	25	(4):	459–474. Linquist,	Stefan,	Edouard	Machery,	Paul	E	Griffiths,	and	Karola	Stotz.	2011.	"Exploring	the	Folkbiological Conception	of	Human	Nature."	Philosophical	Transactions	of	the	Royal	Society	B	366:	444–53. Machery,	Edouard.	2008.	"A	Plea	for	Human	Nature."	Philosophical	Psychology	21	(3):	321–29. Mameli,	Matteo,	and	Patrick	Bateson.	2011.	"An	Evaluation	of	the	Concept	of	Innateness."	Philosophical Transactions	of	the	Royal	Society	of	London	B:	Biological	Sciences	366	(1563):	436–43. Mameli,	Matteo,	and	Patrick	P	G	Bateson.	2006.	"Innateness	and	the	Sciences."	Biology	and	Philosophy	21 (2):	155–88. Medin,	Douglas,	and	Scott	Atran.	2004.	"The	Native	Mind:	Biological	Categorization	and	Reasoning	in Development	and	across	Cultures."	Psychological	Review	111:	960–83. Medin,	Douglas	L.,	and	Scott	Atran.	1999.	"Introduction."	In	Folkbiology,	edited	by	Douglas	L.	Medin	and Scott	Atran,	1–15.	Cambridge,	Mass.:	MIT	Press. Medin,	Douglas,	and	Anthony	Ortony.	1989.	"Psychological	Essentialism."	In	Similarity	and	Anological Reasoning,	edited	by	Stella	Vosniadou	and	Andrew	Ortony,	175–95.	Cambridge:	Cambridge University	Press. Mitchell,	Sandra	D.	2003.	Biological	Complexity	and	Integrative	Pluralism.	Cambridge:	Cambridge University	Press. ---.	2009.	Unsimple	Truths:	Science,	Complexity,	and	Policy.	Chicago:	University	of	Chicago	Press. Narvaez,	D.,	T.	Gleason,	L.	J.	Wang,	and	J.	Brooks.	2013.	"The	Evolved	Development	Niche:	Longitudinal Effects	of	Caregiving	Practices	on	Early	Childhood	Psychosocial	Development."	Early	Childhood Research	Quarterly	28	(4):	759–73. Oyama,	Susan.	2000.	Evolution's	Eye: A	Systems	View	of	the	Biology-Culture	Divide.	Durham,	North Carolina:	Duke	University	Press. Parrott,	R.,	and	R.	A.	Smith.	2014.	"Defining	Genes	Using	'Blueprint'	Versus	'Instruction'	Metaphors: Effects	for	Genetic	Determinism,	Response	Efficacy,	and	Perceived	Control."	Health Communication	29	(2):	137–46. Ramsey,	Grant.	2012.	"How	Human	Nature	Can	Inform	Human	Enhancement:	A	Commentary	on	Tim Lewens's	Human	Nature:	The	Very	Idea."	Philosophy	&	Technology	25	(4):	479–83. doi:10.1007/s13347-012-0087-2. ---.	2013.	"Human	Nature	in	a	Post-Essentialist	World."	Philosophy	of	Science	80	(5):	983–93. ---.	In	Press.	Trait	bin	and	trait	cluster	accounts	of	human	nature.	In:	Elizabeth	Hannon	and	Tim Lewens	(eds)	Why	we	disagree	about	Human	Nature.	Oxford:	Oxford	University	Press. Sterelny,	Kim.	2003.	Thought	in	a	Hostile	World:	The	Evolution	of	Human	Cognition.	Oxford:	Blackwells. ---.	2012.	The	Evolved	Apprentice.	MIT	Press. Sterelny,	Kim,	and	Paul	E	Griffiths.	1999.	Sex	and	Death:	An	Introduction	to	the	Philosophy	of	Biology. Chicago:	University	of	Chicago	Press. Stotz,	Karola.	2010.	"Human	Nature	and	Cognitive-Developmental	Niche	Construction."	Phenomenology and	the	Cognitive	Sciences	9	(4). ---.	2014.	"Extended	Evolutionary	Psychology:	The	Importance	of	Transgenerational	Developmental Plasticity."	Frontiers	in	Psychology	5	(August).	doi:10.3389/fpsyg.2014.00908. ---.	Under	Review.	"Why	Developmental	Niche	Construction	is	not	Selective	Niche	Construction	– and	why	it	matters".	To	appear	in	Interface	Focus,	Special	Issue	"New	Trends	in	evolutionary Biology" Stotz,	Karola,	and	Colin	Allen.	2012.	"From	Cell-Surface	Receptors	to	Higher	Learning:	A	Whole	World	of Experience."	In	Philosophy	of	Behavioural	Biology,	edited	by	Katie	Plaisance	and	Thomas	Reydon, 85–123.	Boston,	MA:	Springer. Stotz,	Karola,	and	Paul	E.	Griffiths.	2008.	"Biohumanities:	Rethinking	the	Relationship	between Biosciences,	Philosophy	and	History	of	Science,	and	Society."	Quarterly	Review	of	Biology	83	(1): 37–45. Stotz,	Karola,	and	Paul	E.	Griffiths.	2016.	"A	Niche	for	the	Genome."	Biology	&	Philosophy	31	(1):	143–57. Sultan,	Sonia	E.	2015.	Organism	and	Environment:	Ecological	Development,	Niche	Construction,	and Adaptation.	Oxford	University	Press. Trevathan,	Wenda.	2011.	Human	Birth:	An	Evolutionary	Perspective.	New	York:	Aldine	de	Gruyter. Waddington,	Conrad	H.	1942.	"Canalisation	of	Development	and	the	Inheritance	of	Acquired	Characters." Nature	150:	563–65. West,	Meredith	J,	and	Andrew	P	King.	1987.	"Settling	Nature	and	Nurture	into	an	Ontogenetic	Niche." Developmental	Psychobiology	20:	549–62. West,	Meredith	J.,	and	Andrew	P.	King.	2008.	"Deconstructing	Innate	Illusions:	Reflections	on	NatureNurture-Niche	From	an	Unlikely	Source."	Philosophical	Psychology	21	(3):	383–95.