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Evolutionary Significance
of tb1 locus in Maize
Aditi Sarkar
L-2019-A-82-M
1
Evolution and Domestication
Evolution
•Change in heritable
characteristics of biological
populations over time
Domestication
•Process of human
interference to direct crop
evolution
2
Stages of Domestication and Diversification
Onset of domestication
Increase in frequency of desirable alleles
Formation of cultivated populations that are
adapted to new environments and local
preferences
Deliberate breeding
3
Domestication
Syndrome
Properties that
distinguish a certain crop
from its wild progenitor
• Larger fruits or grains
• Increase in apical
dominance
• Reduced shattering
4
Maize (Zea mays L.)
3rd staple food crop
Poaceae family
Genus- Zea, group of annual and
perennial grasses
Monoecious, protandrous
5
Geographical origin of maize
• According to Vavilov, origin of maize
is South Central Mexico
• It was supposed to be first
domesticated in the Tehuaccan valley
of Mexico and the original wild form
has been extinct since many years
• Archaeological remains of earliest
maize cob, found at Guila Naquitz
Cave in Oxaca valley of Mexico
(6250 years ago).
6
Evolution of Maize: Three Hypothesis
Tripartite Hypothesis
Recombination Hypothesis
Teosinte Hypothesis
7
8
• Argument
Differences in
chromosome
number and
genetic
constitution
between Zea
and Tripsacum
9
Recombination Hypothesis (Eubanks,1995)
Tripsacum
dactyloides
Zea
diploperennis
Zea
mays
Putative Hybrids
Tripsacorn and
Sundance
10
Teosinte Hypothesis
(Beadle 1939)
• Teosinte cultivated by ancient
people.
• Mutations arose and were selected
• 5 major mutations
• Over the course of time, additional
major plus minor mutations
selected
11
What is teosinte?
• Teosinte is a wild grass in the
Poaceae family that includes
the species Zea mays (in
addition to other wild grass
species: Z.diploperennis, Z.
perennis, and Z. luxurians)
• Z. mays encompasses several
subspecies among which Zea
mays ssp. parviglumis and ssp.
mexicana share a close genetic
relationship with maize.
12
Why teosinte and not tripsacum?
• Teosinte is having same chromosome number and nearly similar
genetic constitution as that of maize
• Similar in morphology to maize except that its branching type
• Tripsacum differs in both chromosome number and genetic
constitution.
• It is grassy type
13
Difference between
maize and teosinte
• Difference in
plant architecture
• Difference in
inflorescence
14
15
Evolution and Domestication of Maize
• The origin of maize was a mystery
• George Beadle suggested that teosinte is the ancestor of maize when
he observed that teosinte and maize have nearly identical
chromosomes
• But many botanists doubted Dr. Beadle’s conclusion because maize
and teosinte have many physical differences.
• Molecular analyses identified one form of teosinte (Zea mays ssp.
parviglumis) as the progenitor of maize
16
Contd..,
• This teosinte grows commonly as a wild plant in the valleys of Southern Mexico
• It is also know as Balsas teosinte
• The domestication of crop plants has often involved an increase in apical
dominance
• Maize exhibits a striking example of this phenomenon, there is a profound
increase in apical dominance compared with teosinte.
• Differences in the morphologies of maize and teosinte were the result of human
selection during the domestication process
17
Cytogenetics and molecular evidence to
evolution
Chromosome number of cultivated maize (Zea mays ssp. mays)
2n = 2x = 20
Chromosome number of teosinte (Zea mays ssp. mexicana)
2n = 2x = 20
Chromosome number of tripsacum (Tripsacum. dactyloides)
2n = 4x = 36
• Mexicana- complete chromosomal pairing and full fertility
• Parviglumis- similar isozyme allelic constitution
18
Genetic evidence of maize domestication
19
Experiment conducted by Dr Beadle in
the 1970s
Teosinte × Maize F1 F2
Prediction of number of genes
Genetic evidence of maize domestication
(Contd..)
• From these data and his mathematical model, he
concluded that four or five genes were responsible for
the differences between teosinte and maize
• In the 1990s, Dr. Doebley and Dr. Stec identified five
genetic regions, that together account for most of the
variation between maize and teosinte, further
supporting Dr. Beadle’s hypothesis.
20
Key domestication genes cloned in maize
Gene Phenotype
tb1 Plant architecture
gt1 Plant architecture
tru1 Plant architecture
tga1 Hardened fruit case
21
Key domestication genes cloned in maize (Contd..)
Gene Phenotype
ZmSWEET4c Seed filling
UB3 Kernel row number
ZmSh1-1 Shattering
ra1 Inflorescence architecture
22
Teosinte glume architecture (tga1) and teosinte branched one (tb1)
• regulatory genes
• transformation (Teosinte  maize)
• tga1  expression of the traits associated with the seeds.
• tb1  expression of the traits associated with branching and
inflorescence.
• Difference in expression pattern between teosinte and maize
version
23
24
Teosinte branched 1 (tb1)
was identified as a major QTL involved in apical dominance
•is a member of the TCP family of transcriptional regulators
•rice homolog of tb1- OsTB1
25
Contd..,
tb1 also targets other domestication loci in maize
involved in regulation of some phytohormones
also modulates FT1 activity
26
Contd..,
tbl-ref - recessive, null or loss of function mutant that
produces plants with long lateral branches tipped by tassels
at upper nodes and tillers at basal nodes
Tb1+maize - dominant maize allele, produces short
lateral branches tipped by ears at upper nodes and few or
no tillers at basal nodes
tb1+ teosinte – weak function, a partial teosinte
branched phenotype
27
A Model for tb1 (Doebley et al 1995)
• Plants respond to
environmental conditions by
altering their architecture.
• Teosinte also exhibits a
plastic response to local
environment
• tb1 is involved in regulating
the plasticity response
28
A Model for tb1 (Contd..,)
Favorable environmental conditions
Tb1+ teosinte turned off
Axillary meristems develop
fully into tillers or long lateral
branches tipped by tassels
29
A Model for tb1 (Contd..)
Unfavorable environmental conditions
Tb1+ teosinte turned on
Few or no tillers and only
short lateral branches
tipped by ears
30
The Role of tb1 in Maize Evolution
In teosinte, tb1 should be off or expressed at low levels in the
primordia that form the primary branches
tb1 should also be off (or at low levels) in the inflorescence
primordia, so that tassels would be formed rather than an ear
Thus, the evolution of maize required an increase in tb1
expression
31
Data supporting the tb1 model
Maize allele of tb1 is expressed at about twice the level of teosinte allele in
immature axillary branches and the inflorescence primordia
In-situ hybridization in teosinte showed no sign of tb1 expression
in axillary buds, where maize shows strong expression
No fixed amino acid differences between maize and teosinte
32
Quantification of tb1 mRNA levels
33
Conclusion
• Present races of maize was achieved through selection for
overexpression of the teosinte branched1 (tb1) transcription factor.
• Based on the study of genomic changes during maize domestication, it
was found that many traits are affected by limited number of genes.
• Thus, it is possible that we could improve specific maize traits by
manipulating only a handful of genes to meet modern demands of
diverse maize types.
34
THANK YOU
35

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Evolutionary significance of tb1 locus in Maize

  • 1. Evolutionary Significance of tb1 locus in Maize Aditi Sarkar L-2019-A-82-M 1
  • 2. Evolution and Domestication Evolution •Change in heritable characteristics of biological populations over time Domestication •Process of human interference to direct crop evolution 2
  • 3. Stages of Domestication and Diversification Onset of domestication Increase in frequency of desirable alleles Formation of cultivated populations that are adapted to new environments and local preferences Deliberate breeding 3
  • 4. Domestication Syndrome Properties that distinguish a certain crop from its wild progenitor • Larger fruits or grains • Increase in apical dominance • Reduced shattering 4
  • 5. Maize (Zea mays L.) 3rd staple food crop Poaceae family Genus- Zea, group of annual and perennial grasses Monoecious, protandrous 5
  • 6. Geographical origin of maize • According to Vavilov, origin of maize is South Central Mexico • It was supposed to be first domesticated in the Tehuaccan valley of Mexico and the original wild form has been extinct since many years • Archaeological remains of earliest maize cob, found at Guila Naquitz Cave in Oxaca valley of Mexico (6250 years ago). 6
  • 7. Evolution of Maize: Three Hypothesis Tripartite Hypothesis Recombination Hypothesis Teosinte Hypothesis 7
  • 8. 8
  • 9. • Argument Differences in chromosome number and genetic constitution between Zea and Tripsacum 9
  • 11. Teosinte Hypothesis (Beadle 1939) • Teosinte cultivated by ancient people. • Mutations arose and were selected • 5 major mutations • Over the course of time, additional major plus minor mutations selected 11
  • 12. What is teosinte? • Teosinte is a wild grass in the Poaceae family that includes the species Zea mays (in addition to other wild grass species: Z.diploperennis, Z. perennis, and Z. luxurians) • Z. mays encompasses several subspecies among which Zea mays ssp. parviglumis and ssp. mexicana share a close genetic relationship with maize. 12
  • 13. Why teosinte and not tripsacum? • Teosinte is having same chromosome number and nearly similar genetic constitution as that of maize • Similar in morphology to maize except that its branching type • Tripsacum differs in both chromosome number and genetic constitution. • It is grassy type 13
  • 14. Difference between maize and teosinte • Difference in plant architecture • Difference in inflorescence 14
  • 15. 15
  • 16. Evolution and Domestication of Maize • The origin of maize was a mystery • George Beadle suggested that teosinte is the ancestor of maize when he observed that teosinte and maize have nearly identical chromosomes • But many botanists doubted Dr. Beadle’s conclusion because maize and teosinte have many physical differences. • Molecular analyses identified one form of teosinte (Zea mays ssp. parviglumis) as the progenitor of maize 16
  • 17. Contd.., • This teosinte grows commonly as a wild plant in the valleys of Southern Mexico • It is also know as Balsas teosinte • The domestication of crop plants has often involved an increase in apical dominance • Maize exhibits a striking example of this phenomenon, there is a profound increase in apical dominance compared with teosinte. • Differences in the morphologies of maize and teosinte were the result of human selection during the domestication process 17
  • 18. Cytogenetics and molecular evidence to evolution Chromosome number of cultivated maize (Zea mays ssp. mays) 2n = 2x = 20 Chromosome number of teosinte (Zea mays ssp. mexicana) 2n = 2x = 20 Chromosome number of tripsacum (Tripsacum. dactyloides) 2n = 4x = 36 • Mexicana- complete chromosomal pairing and full fertility • Parviglumis- similar isozyme allelic constitution 18
  • 19. Genetic evidence of maize domestication 19 Experiment conducted by Dr Beadle in the 1970s Teosinte × Maize F1 F2 Prediction of number of genes
  • 20. Genetic evidence of maize domestication (Contd..) • From these data and his mathematical model, he concluded that four or five genes were responsible for the differences between teosinte and maize • In the 1990s, Dr. Doebley and Dr. Stec identified five genetic regions, that together account for most of the variation between maize and teosinte, further supporting Dr. Beadle’s hypothesis. 20
  • 21. Key domestication genes cloned in maize Gene Phenotype tb1 Plant architecture gt1 Plant architecture tru1 Plant architecture tga1 Hardened fruit case 21
  • 22. Key domestication genes cloned in maize (Contd..) Gene Phenotype ZmSWEET4c Seed filling UB3 Kernel row number ZmSh1-1 Shattering ra1 Inflorescence architecture 22
  • 23. Teosinte glume architecture (tga1) and teosinte branched one (tb1) • regulatory genes • transformation (Teosinte  maize) • tga1  expression of the traits associated with the seeds. • tb1  expression of the traits associated with branching and inflorescence. • Difference in expression pattern between teosinte and maize version 23
  • 24. 24
  • 25. Teosinte branched 1 (tb1) was identified as a major QTL involved in apical dominance •is a member of the TCP family of transcriptional regulators •rice homolog of tb1- OsTB1 25
  • 26. Contd.., tb1 also targets other domestication loci in maize involved in regulation of some phytohormones also modulates FT1 activity 26
  • 27. Contd.., tbl-ref - recessive, null or loss of function mutant that produces plants with long lateral branches tipped by tassels at upper nodes and tillers at basal nodes Tb1+maize - dominant maize allele, produces short lateral branches tipped by ears at upper nodes and few or no tillers at basal nodes tb1+ teosinte – weak function, a partial teosinte branched phenotype 27
  • 28. A Model for tb1 (Doebley et al 1995) • Plants respond to environmental conditions by altering their architecture. • Teosinte also exhibits a plastic response to local environment • tb1 is involved in regulating the plasticity response 28
  • 29. A Model for tb1 (Contd..,) Favorable environmental conditions Tb1+ teosinte turned off Axillary meristems develop fully into tillers or long lateral branches tipped by tassels 29
  • 30. A Model for tb1 (Contd..) Unfavorable environmental conditions Tb1+ teosinte turned on Few or no tillers and only short lateral branches tipped by ears 30
  • 31. The Role of tb1 in Maize Evolution In teosinte, tb1 should be off or expressed at low levels in the primordia that form the primary branches tb1 should also be off (or at low levels) in the inflorescence primordia, so that tassels would be formed rather than an ear Thus, the evolution of maize required an increase in tb1 expression 31
  • 32. Data supporting the tb1 model Maize allele of tb1 is expressed at about twice the level of teosinte allele in immature axillary branches and the inflorescence primordia In-situ hybridization in teosinte showed no sign of tb1 expression in axillary buds, where maize shows strong expression No fixed amino acid differences between maize and teosinte 32
  • 33. Quantification of tb1 mRNA levels 33
  • 34. Conclusion • Present races of maize was achieved through selection for overexpression of the teosinte branched1 (tb1) transcription factor. • Based on the study of genomic changes during maize domestication, it was found that many traits are affected by limited number of genes. • Thus, it is possible that we could improve specific maize traits by manipulating only a handful of genes to meet modern demands of diverse maize types. 34

Editor's Notes

  1. Why teosinte has been accepted and not tripsacum. Maize is strongly allied with teosinte in terms of chromosome morphology and number.
  2. Teosinte is highly branched; its bushy form has many stems (called “tillers”) and produces heads with two rows of five to 12 seeds at the top of each stalk. By contrast, a maize plant usually has just one central stalk that produces a few ears, each with hundreds of kernels in six to 18 rows. Another stark difference is that teosinte’s seeds, or kernels, are surrounded by a hard fruitcase. This protective covering enables seeds to survive the digestive tracts of birds and grazing animals. When the seeds are excreted with animal waste, they can germinate, effectively using the animals as dispersal agents to spread the plants to distant locales. In contrast, the fruitcase of maize is greatly reduced and develops into part of the cob. This leaves the kernel exposed or naked and thus easily digested by animals.
  3. Apical dominance- the concentration of resources in the main stem of the plant and a corresponding suppression of axillary branches).
  4. Gt1= grassy tillers 1 tru1= tassels replace upper ears1 Tga1= teosinte glume architecture1
  5. ZmSWEET= sugar transporter ZmSh1 = shattering genes UB3 = unbranched 3 ra1 = ramose 1
  6. In teosinte tb1 is functional and is normally expressed in the secondary axillary meristem where it controls their conversion into ear shoots. tb1 is not normally expressed in the primary axillary meristems of teosinte so that THESE ARE ABLE TO develop into elongated tassel-tipped branches. During the domestication of maize, humans selected an allelic variant of tb1 that is expressed in primary axillary meristems (and probably has a high level of expression) such that these form ear shoots rather than elongated tassel-tipped branches.
  7. .
  8. unfavorable environmental conditions= such as a high level of competition from surrounding vegetation, shading, and restricted moisture) by growing into slender unbranched plants (strong apical dominance), or correspondingly they can respond to favorable local environmental conditions by growing into robust highly branched plants (weak apical dominance). tbl is involved in regulating this response by specifying the fate of axillary meristems
  9. Under favorable environmental conditions, tbl+ teosinte is turned off, allowing axillary meristems to develop fully into tillers or long lateral branches tipped by tassels
  10. Under unfavorable conditions, tbl+ teosinte is turned on so that the plant produces few or no tillers and only short lateral branches tipped by ears. Thus, tbl is hypothesized to be a locus involved in the plastic response of the teosinte plant to its local environment by altering plant architecture.
  11. This would enable the growth of these primordia into fully elongated branches………..tb1 should also be off in the inflorescence primordium terminating the primary branch and its stamen primordia so that tassels would not be repressed. Increase in tb1 exp in prim ax branch prim and its terminal inflorescence, so that they form ears rather than tassel tipped branches.
  12. No amino acid diff shows that change in protein function has not occurred, instead change in expression level is involved
  13. Expression is higher in maize allele as compared to teosinte allele
  14. Taking the waxy endosperm trait as an example, mutation of only one gene (Wx) leads to an amylopectin content of nearly 100% in the endosperm