3. • Neolithic revolution/Agriculture
revolution: 12000 to 10000 years
ago
• Changes:
– The transition from hunting-
gathering to plant agriculture
– The formation of villages
– One of the key technological
elements of the transition to
agriculture was DOMESTICATION.
Introduction
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Plant domestication is the genetic
modification of a wild species to create a
new form of a plant altered to meet human
needs(Doebley et al., 2006)
The process by which humans actively
interfere with and direct crop evolution.
What is domestication..?
Continuum of increasing codependence
between plants and people
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Incidental- Didn’t happen on purpose. Hunter/gatherers dropped
seeds, scared off natural herbivores, disrupted natural environments
so that plants could grow.
Directed- Humans and plants became dependent on each other, so
better plants helped people get healthier, planting more (and maybe
improved) plants, etc.
Types of Domestication
Agriculture- Human intervention in crop
Husbandry, Cultivation and Selection.
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Where did it happen….??
(Balter, 2007)
Africa
Meso-America
Near East China
South-America
South-East Asia
Gepts, P., 2004
6
7. 16-Dec-17 PG seminar 7
How did it happen…???
Is Artificial Selection
Analogous to Natural
Selection?
Unconscious selection: ‘‘that which
follows from men naturally preserving
the most valued and destroying the less
valued individuals, without any thought
of altering the breed.’’ (Darwin, 1868)
It was no different from natural selection
Humans change the conditions in which cultivated species live
and reproduce,
The phenotypic changes associated with domestication are
likely to have arisen via unconscious selection
(Harlan, 1992)
Natural
selection
Domestication
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What made domesticates to differed from wild….??
• There is a common suite of traits that
distinguishes most seed and fruit crops
from their progenitors (Hammer, 1984)
These distinct suite of traits later termed the
“domestication syndrome” would likely be
selected for during the initial stages of
domestication (Harlan et al., 1973)
Domestication syndrome:
It is the subset of traits that collectively
form the morphological and
physiological differences between crops
and their wild progenitors.
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‘Domestication Syndrome’ traits
A variety of physiological
changes are also involved.
a loss of seed dormancy,
a decrease in bitter
substances in edible
structures
changes in photoperiod
sensitivity
synchronized flowering
Compared to their progenitors,
food crops typically have
larger fruits or grains,
more robust plants
more determinate growth or
increased apical dominance
a loss of natural seed dispersal
often have fewer (although
larger) fruits or grains per plant
than their progenitors.
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Diversification genes ..
‘Diversification’ genes are the target of selection for phenotypic variation
among varieties of a crop, such as different types of starch or flavor
Glutinous or sticky phenotype of cooked cereal grains, reflecting the
absence (or near absence) of the starch amylose in the endosperm - the
glutinous phenotype is favored in select varieties of rice (primarily a subset
of japonica varieties of O. sativa), maize and foxtail millet, and is
controlled by unique mutations at the Waxy gene in all these crops.
Rice BADH2 gene (2-Acetyi 1-Pyrrlione) - the aromatic phenotype has also
been generated via a variety of mutations at the same gene
(Gross and Olsen, 2010)
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Methods for identifying domestication genes
Biparental QTL mapping
Association Mapping Using Unrelated Individuals
QTL Mapping Using Advanced Intercross
Populations
Genomic scans
Genome Resequencing and Screening for
Selection Signatures
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Classical Examples…
Teosinte branched 1 (tb1) QTL of maize controls the difference in apical dominance in maize
and teosinte
tb1, it acts as transcriptional regulators, a class of genes involved in the transcriptional
regulation of cell cycle
Teosinte glume architecture1 (tga1) was identified as a QTL controlling the formation of the
casing that surrounds the kernels of the maize ancestor, teosinte
tga1 is a member of the squamosa-promoter binding protein (SBP) family of transcriptional
regulators
Fruitweight2.2 (fw2.2) was identified as a large effect QTL controlling 30% of the difference
in fruit mass between wild and cultivated tomato
fw2.2 acts as a negative regulator of cell division in the fruit, perhaps via some role in cell-to-
cell communication
Q is a major gene involved in wheat domestication that affects a suite of traits, including
The tendency of the spike (ear) to shatter,
The tenacity of the chaff surrounding the grain, and
The spike is elongated as in wild wheat or compact like the cultivated forms
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shattering4 (sh4) is a major QTL controlling whether the seed fall off the plant
(shatter) as in wild rice or adhere to the plant as in cultivated rice
sh4 encodes a gene with homology to Myb3 transcription factors.
A single amino acid change in the predicted DNA binding domain converts plants
from shattering to non-shattering
Rc is a domestication-related gene required for red pericarp in rice
Two independent genetic stocks of Rc revealed that the dominant red allele differed
from the recessive white allele by a 14-bp deletion within exon 6 - originated in
japonica cultivar and spread into indica cultivars.
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Super-domestication
The processes that lead to a domesticate with dramatically increased yield that
could not be selected in natural environments without new technologies.
The array of genome manipulations enable barriers to gene exchange to be
overcome and have lead to super-domesticates with
– dramatically increased yields,
– resistances to biotic and abiotic stresses, and with
– new characters for the marketplace.
Hybrid rice can be considered a super-domesticate
Conversion of a crop from C3 to C4 photosynthesis
would certainly be a super-domesticate.
(Vaughan et al. 2007)
PLANT
BREEDERS
GENOMIC
SCIENTISTS
SUPER
DOMESTICATION
leads to
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Effect of selection during domestication
Selection is expected to reduce diversity at domestication related genes and
tightly linked loci
One common feature of the domesticated genomes is the reduction of
genetic diversity in crops relative to the wild progenitors
This reduction has two causes:
Genetic bottleneck
Selective sweep
(Tang et al., 2010)
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It depends on
The bottleneck population size
The duration of bottleneck
The loss in diversity was not experienced equally by all genes in the genome
(Tang et al., 2010)
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Effect of Artificial Selection on Genetic Diversity of
Maize Genes.
Bottleneck
effect
(Yamasaki et al., 2005)
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A selective sweep is the reduction or elimination of variation among
the nucleotides in neighboring DNA of a mutation as the result of recent and
strong positive natural selection
A strong selective sweep results in a region of the genome where the
positively selected haplotype (the mutated allele and its neighbours) is
essentially the only one that exists in the population, resulting in a large
reduction of the total genetic variation in that chromosome region.
Selective sweep
Size of
selective
sweep
Time of selection
Strength of selection
Recombination rates
(Tang et al., 2010)
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Case study:1
Aim of the study:
• Attempt identify gene responsible for prostrate growth
and erect growth habit
• Mapping the gene on to chromosome
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Linkage analysis : F2 derived from cross between YIL 18 and Teqing showed that
prostrate growth was completely controlled by a single semi-dominant gene, PROG1
(PROSTATE GROWTH 1), located between SSR markers RM298 and RM481 on
short arm of chromosome 7
Output of the study
Positional cloning study: demonstrate that in the O. rufipogon genome was a
key gene (PROG1) controlling prostrate growth.
Tissue specificity study: they followed GUS reporter gene method
• detected GUS expression in the tiller base, leaf-sheath pulvinus and lamina
joint, but not in the root, leaf blade and culm
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Case study:2
Preamble:
Why did a crop domestication gene hinder breeding with a
modern breeding gene responsible for the beneficial ‘‘jointless’’ trait
in tomato, and how can this genetic interaction be overcome and
exploited?
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Understanding crop domestication- Implications
Up- and down-regulation of transcription factors
Application of selection screens to identify genes contributing
to the success of best varieties
QTL cloning for key agronomic genes
Screening wild relatives and unimproved varieties to recover
superior alleles (allele mining) that failed to pass through the
domestication and improvement bottlenecks
Since their origin, hunting and gathering had been the primary mode of subsistence for modern humans. But around 12,000 years ago, humans switched from a hunter-gatherer lifestyle to an agricultural lifestyle. This transition in human behavioural ecology is known as ‘‘the Neolithic revolution’’. The Neolithic revolution has marked one of the most profound changes in human evolution.
Domestication can be described as a set of consecutive stages that begins with the onset of domestication followed by an increase in the frequency of a set of desirable traits (the domestication traits), and which culminates with the emergence of cultivated populations adapted to both human needs and a cultivated environment.
this change exerts selection on the population even in the absence of a choice or predetermined goal by the cultivator.
Oasis: Symbiotic association between human animal and plants due to gathering around few oasis and water resources.
Hilly flank: Agriculture began in upland location with frequent rainfall so that crop can grown Without supplemented irrigation.
Population pressure: Population increases in near East area/ South West Asia upset the balance between People and food , forcing people to turn to an agricultural way of life .
Maize
Wild
Multiple stalks and long branches
Teosinte ear has its grains enclosed in the triangular casing that comprises the ear
cultivated
Single stalk
Maize ear bears its grain naked on the surface of the ear
Rice
Wild rice has a panicle that shatters…………….Cultivated rice has a solid panicle of grains
challenging task is to define a domestication syndrome, which is the subset of traits that collectively form the morphological and physiological differences between crops and their wild progenitors.
Transforming plant breeding into crop engineering to meet the human need for increased crop yield with the minimum environmental impact. We consider this to be ‘super-domestication
Early farmers used only a limited number of individuals of the progenitor species, much of the genetic diversity in the progenitor was left behind
A population bottleneck (or genetic bottleneck) is an evolutionary event in which a significant percentage of a population or species is killed or otherwise prevented from reproducing
This evidence suggests that mutation of a single gene can improve plant architecture and yield in crop domestication;