Expert answer:Compare and contrast the relationship of environme

Expert answer:Paper due 5 PM FRIDAY, Dec. 8. Please upload your paper to the turnitin.com icon on TritonEd AND submit a paper copy before FRIDAY, Dec. 8. (either in class or in the Drop Box on the 2nd floor of the Social Sciences Building to the right of the Anthropology Department office). Clearly mark the course number and your section on the front of your exam. If you would like your paper to be returned in the basket in the same place next quarter, please sign and attach the included Buckley Waiver. in no more than six (6) pages. Use appropriate course readings plus AT LEAST TWO ADDITIONAL SCHOLARLY SOURCES to build your argument. Discuss SPECIFIC ARCHAEOLOGICAL EXAMPLES in detail to support your arguments. Text must be 12 point, double spaced, with 1 inch margins (not including bibliography). Cite sources for all your important points, whether paraphrased or direct quotations. Indicate which question you are answering.1) States developed independently in very different environmental settings. Environmental factors can both stimuli or constraints to the development and sustainability of ancient states.Compare and contrast the relationship of environmental factors to the rise and fall of complex civilizations in at least two civilization areas. How do climate, geography, geology, plant and animal species, agricultural potential, resource abundance or scarcity affect the development of states in each region?
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Evolutionary Anthropology 15:105–117 (2006)
ARTICLES
Central Questions in the Domestication of Plants
and Animals
MELINDA A. ZEDER
Along with symbolic communication, tool use, and bipedalism, the domestication of plants and animals, together with the associated emergence of agriculture,
stands as one of the pivotal thresholds in human evolution. For more than a
hundred years researchers have wrestled with the questions of what domestication is, how it is detected, and why it happened. The past decade in particular has
witnessed a remarkable acceleration of interest in domestication, thanks to advances in our ability to detect the context, timing, and process of domestication in
a wide array of different plant and animal species around the world.1 This review
focuses on overarching issues of defining, documenting, and explaining the domestication of plants and animals, tracing a path through often discordant viewpoints to offer some new perspectives.
DEFINING DOMESTICATION
All approaches to defining domestication in both plants and animals recognize that domestication involves a
relationship between humans and target plant or animal populations. There
are, however, distinct and often discordant perspectives taken regarding
the balance of power in this relationship and its central defining features
Melinda A. Zeder is Director of the Archaeobiology Program of the Smithsonian Institution’s National Museum of Natural History. Her research focuses on
questions of domestication, origins of agriculture, and the environmental and social impacts of early agricultural economies in the ancient Near East. She is the
lead editor of the volume, Documenting
Domestication: New Genetic and Archaeological Paradigms, with co-editors Eve
Emshwiller, Daniel G. Bradley, and Bruce
D. Smith, published in spring of 2006 by
the University of California Press.
Key words: domestication; origins of agriculture;
plants; animals; climate change; demographic
pressure; social forces; Southwest Asia
© 2006 Wiley-Liss, Inc.
DOI 10.1002/evan.20101
Published online in Wiley InterScience
(www.interscience.wiley.com).
(Fig. 1). Many approaches to defining
domestication, especially those focusing on animals, emphasize the dominant role humans play in assuming
“mastery” over all aspects of the reproduction, movement, distribution,
nourishment, and protection of domesticates.2–5 Integral to definitions
that place humans in control of the
process is the notion of intentionality,
that humans with foresight and deliberate intent intervened in the life cycle
of target plant and animal populations and assumed responsibility for
their care to meet specific and welldefined objectives serving human
needs. Also often associated with this
emphasis on the human dimension is
the notion that domestication involves a fundamental change in socioeconomic organization in which successive generations of domesticates
become integrated into human societies as objects of ownership.3,6
Other researchers object to “anthropocentric” approaches to defining domestication that portray domesticates
as passive pawns in the process, pointing out that domesticates also reap
benefits through vastly enhanced reproductive fitness and expanded ranges.7 Those operating within an evolutionary biology perspective, in
particular, maintain that the relationship between humans and domesticates is no different from other mutualistic relationships in the “natural
world” that bring together species like
ants and aphids in partnerships of increasing co-dependency.8 Moreover,
as one moves further along the spectrum, from a relatively balanced mutualistic perspective to ones that focus
on the domesticate, the role of deliberate human intent declines. The
more extreme positions at this end of
the spectrum tip the balance in favor
of the domesticate, which is seen as
manipulating its human partners for
its own evolutionary advantage, ensnaring humans in a relationship that
may have actually reduced human fitness.9
Another axis of variation in definitional approaches to domestication is
the relative primary given to genetic
and associated morphological change.
An emphasis on genetic change and
its phenotypic expression is particularly common among researchers focusing on plant domestication, especially the domestication of largeseeded annuals, where human
intervention results in fairly rapid genetic changes with easily observed
phenotypic expressions.9,10 Some researchers focusing on animals also
see genetic isolation and subsequent
quick-onset morphological change as
essential attributes of domestication.11
The requirement that domesticates
show evidence of morphological or
even genetic change, however, is not
universally accepted. Nor is the basic
premise underlying this requirement:
that the process of domestication is
contingent on reproductive isolation
and resultant genetically driven mor-
106 Zeder
ARTICLES
Figure 1. Definitions of domestication tend to fall somewhere along three axes of variation. Definitions that award the balance of power
in the domestic relationship to humans tend to stress human intentionality and the social and economic impacts of domestication.
Definitions that tip the balance of power in favor of the domesticate tend to discount the role of human intentionality in the process and
stress its biological impacts on the domesticate.
phological change. This is particularly
true for animals, where morphological change, when it occurs at all, is
often both delayed and difficult to tie
directly to domestication.12 As a result, many researchers define animal
domestication not in terms of observed genetic or morphological
change, but in terms of causal human
behavior. According to this view, domestication falls along a continuum
of increasing human intervention
ranging from predation to genetic engineering13 in which there are varying
degrees of investment in altering an
animal’s natural behavior (its movement, breeding schedule, or population structure) to suit human
needs.6,14 A similar view is becoming
increasingly common in considerations of plant domestication especially perennial plants such as root
crops propagated through vegetative
cloning or very long-lived tree crops in
which genetic and morphological
change may be less automatic and
more subtle than in annual seed
crops.15 Smith,16 for example, maintains that for both plants and animals
the central defining feature of domestication and the creation of domesticates is the nature of the “ongoing relationship of intervention initiated
and sustained by humans.” This em-
phasis on the evolving relationship between humans and plant or animal
populations turns attention away
from a range of secondary consequences of domestication, such as genetic and morphological change or social notions of property, and properly
returns it to a consideration of the
new partnership that humans create
with target populations.
Domestication does indeed have
many features in common with other
mutualistic
relationships
among
plants and animals. Both partners in
the relationship of domestication
clearly derive benefits. Plant and animal partners benefit in increased reproductive fitness and range expansion. Human partners gain increased
security and predictability in their access to resources of interest. Both
partners respond to this relationship
in ways that enhance respective payoffs and further deepen their mutual
investment in its continuation. But
the mutualism that lies at the heart of
the domestication process involving
human societies and target plant and
animal populations varies in significant ways from other similar relationships found in nature.
In a recent overview of the homologies between human agriculturists
and fungus-growing ants, Schultz and
coworkers17 highlight the many parallels between these convergent forms
of mutualism while also underscoring
key qualitative differences between attine and human agriculture. The codependent relationships between
farmer ants and domesticated fungi
are the result of a gradual co-evolutionary process based on mutation-induced behavioral and morphological
change in both partners. Humans, on
the other hand, are capable of modifying their behavioral repertories
through “trial and error, observation,
and imitation.”17 That ability enables
humans to rapidly develop behavioral
strategies aimed at meeting consciously recognized needs. The highly
developed human ability for cultural
transmission of learned behavior,
Shultz and coworkers argue, greatly
accelerated the adaptive modification
of human behavior, shifting the balance of power in the emergent mutualism. Humans quickly assume a
dominant role because they are free to
choose among genetic variants in the
partner population, to manipulate the
behavior and life history of symbionts
(even to their own detriment), or to
terminate the relationship with one
partner symbiont and choose another.
This is where intentionality comes
into the picture. It is true that humans
Domestication of Plants and Animals 107
ARTICLES
could not have foreseen the adaptive
responses by plant and animal partners to the new selective factors
brought into play by the relationship
of domestication. Nor did humans
likely appreciate the long-term benefits (or the negative consequences)
that might accrue from domestication
and the subsequent development of
agricultural economies. However, ruling out this kind of prescience on the
part of humans does not take intentionality out of the picture. While they
might not have understood the principles of genetic engineering, humans
could appreciate the fact that tending,
nurturing, and intervening in the life
cycle of certain plants and animals
yielded various immediate benefits.
On the basis of these returns they
could then consciously and deliberately decide to continue to engage in
these behaviors, and to elaborate on
them, instead of engaging in other
strategies.
Intentionality, then, becomes the
key factor that distinguishes domestication from other similar mutualistic
relationships in nature. The deliberate
role humans take in actively pursuing
the domestic partnership also distinguishes it from other biological relationships between humans and plants
and animal species, such as commensal relationships with mice, sparrows,
or weeds that take advantage of new
niches created by human habitation.
Definitions that try to pigeon-hole domestication as either a cultural or biological process are bound to come up
short. Clearly, domestication has a biological component as a mutualistic
relationship between humans and
plant or animal symbionts. Just as
clearly, however, human intentionality sets domestication apart from
other forms of mutualism. The
uniqueness of the relationship comes
from its cultural component and the
dominant role humans play in consciously and deliberately perpetuating
it to their own advantage.
If the process of domestication is
best viewed as a form of mutualism
that is asymmetrically enhanced by
the human ability to culturally transmit learned behavior, then at what
point along this developmental trajectory does the plant or animal partner
become a domesticate? Is there a
threshold that, once crossed, separates the “wild” from the “domestic”?
If so, what does this threshold look
like? To some extent, it remains a
matter of personal preference to decide just when a domestic subsection
of a plant or animal species has been
created. Threshold criteria that require total genetic isolation and emergent speciation or complete dependence on humans for survival set a
very high bar that many, if not most,
widely accepted domesticates would
fail to clear. Even somewhat looser
standards that involve a lesser degree
of genetic modification in the target
plant or animal population, or a certain level of human investment in
propagating, nurturing, or owning the
resource, run the risk of constructing
artificial boundaries along what was
really a more seamless incremental
process.
Ducking the issue by adopting the
term “proto-domesticate” also does
not help much. This term implies that,
if just given enough time and perhaps
a little more investment by either partner, full domestic status would be
achieved. The actual trajectory of domestication, however, is highly contingent on a wide range of factors,
including the ability of the plant or
animal to take advantage of the relationship, the strategies and accompanying technologies humans develop to
manage the resource, and its changing value vis-à-vis available alternative
resources. In some plant and animal
species, genetic modification and
more focused human investment in
the resource may quickly follow. In
others there may be a long and very
stable relationship involving fairly
minimal commitment by either partner. Further, it appears that budding
domestic relationships sometimes fail
altogether, never moving beyond an
initial courtship phase.
It is best to step back and not focus
too closely or obsessively on defining
the exact demarcation between domestic and wild, and to turn, instead,
to a consideration of the full span of
the evolving nature of domestic relationships. Different stages in the evolution of this relationship might be
characterized by the degree of investment by both partners (Fig. 2). For the
plant or animal, this would involve the
extent of genetic modification made in
response to new selective pressures,
the degree of its genetic isolation from
populations not involved in the partnership, the nature of subsequent
morphological or behavioral change,
and its increasing co-dependency on
humans. For humans, this might be
the level of investment in the production of the resource; that is, in tilling,
watering, burning, and land clearance, sowing, and transplanting
plants, or in taming, protecting, herding, culling, and selectively breeding
animals. It might also include the degree of incorporation of domesticates
within the socio-economic organization of the human groups investing in
its production.
By expanding the scope of inquiry
to encompass the vast “middle
ground” between foraging and farming, hunting and herding,16 we can
approach a deeper, more comprehensive, and ultimately more informative
appreciation of the range of possibilities open to humans and their plant
and animal partners. This expanded
territory of investigation includes the
stable, long-lived systems of low-level
food production involving a mix of
both morphologically altered and
nonaltered domesticates, as well as
“wild” resources, featured in recent
books on indigenous resource management in California and the Northwest Coast.18,19 At the other end of the
spectrum are highly structured agricultural economies with complete dependence on domesticates and total
investment in their production. Trying to understand the full richness of
the various ways in which the domestic partnership may manifest itself in
different contexts, we, in turn, stand a
much better chance of being able to
document and explain domestication.
DOCUMENTING
DOMESTICATION
In both plants and animals, this effort requires identifying clear-cut
markers that can be explicitly linked
to a specific aspect or stage of the unfolding domestication process.1,20 Different markers may be more effective
in detecting different stages of this
process. Markers will also vary depending on the biology of the domes-
108 Zeder
ARTICLES
Figure 2. Domestication is best viewed as an evolving of mutualism between humans and populations of plants or animals. The relationship
can be characterized along various scales of investment by either the human or the plant or animal partners. All of these scales usually are
involved in the process of domestication, though they often operate independently of one another. The degree of change along each
scale is contingent on the biology of the species involved, as well as the ecological and cultural circumstances of the human partners.
Attempting to distinguish just where and along what scale domestication occurs is not only difficult, but may not be very useful.
ticate and its relationship with humans. There are, in particular,
fundamental differences in the selective pressures on plants and animals
undergoing domestication, and, as a
result, in the corresponding markers
used to document plant and animal
domestication.
Selective pressures on plants, especially annuals, tend to operate directly
on morphological traits that can, in
turn, be used as unambiguous markers of domestication.15 Morphological
impacts of the domestication of annuals may come about as largely automatic responses to human planting
and harvesting that result in such
changes as increased seed size, thinner seed coats, reconfiguration of seed
head architecture, or the development
of indehiscent seed pods.15,21,22 Intentional selection for specific morphological attributes in annual plants,
such as larger fruit size, appear to
happen later in the developing relationship of domestication.15
Perennial plants sustained by transplanting root fragments, on the other
hand, are not subjected to the same
seed-bed pressure and human harvesting selective pressures that result
in the morphological markers used to
document domestication in annual
seed plants.15 At the same time, however, because there may be more of an
opportunity for humans selectively to
replant root fragments with desired
traits, these plants may respond fairly
quickly to deliberate human selection
in the development of larger fruits, the
loss of chemical defenses against herbivory, or changes in sugars and
starches.23 While many of these crops
were grown in tropical areas with
poor preservation of plant macro-fossils, the development of breakthrough
techniques for the recovery and identification of plant micro-fossils (that
is, phytoliths and starch grains) has
made it possible to detect these domestication-induced morphological
changes in root and other crop plants
(Fig. 3).24,25
Recent years have seen an increase
the use of nonmorphological markers
of the intensification of human-plant
interactions that may precede clearcut evidence of morphological change
in plants. Evidence of land clearance,
modification of natural drainage systems, intentional burning, and
changes in the composition of weedy
plants in archeological assemblages
have all been effectively used to track
human modification of landscapes
and plant communities as part of the
domestication process.26 –28 The occurrence of plant macro- or micro fossils in areas thought to be far outside
their natural range has also been interpreted as evidence of human transport and tending of plants.26,29,30
There are special challenges to finding markers of animal domestication.
This is because the leading-edge pressures on animals undergoing domestication are likely to focus on behav-
Domestication of Plants and Animals 109
ARTICLES
ioral attributes rather than on
morphological traits.31 There are a variety of behaviors that probably made
certain animal species better candidates for domestication; among them
tolerance of penning, a social structure based on dominance hierarchies,
sexual precocity, weak alarm systems
and, above all, reduced wariness and
aggression.32 Behavioral responses to
domestication in animals elaborated
on these initial preselection qualifying
attributes and include a general reduction in responsiveness to environmental stimuli, reduced activity levels, increased social compatibility,
and intensified sexual behavior.4,33
Many morphological traits commonly seen in domestic animals are
thought to be linked to these behavioral changes. These attributes include piebald coats, lop ears and, of
special importance here, reduced
brain size and an overall juvenilization of cranial form.4,33 This latter feature may result in a shortened muzzle,
tooth crowding, and reduction in
tooth size, trait …
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