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Arabidopsis thaliana

Model organism for genetic


analysis in developmental plant
biology
Relatively short generation time
About 2 months
Small genome size similar to

Drosophila and C. elegans


14 x 107 bp

Can be subjected to mutagens to

generate mutations altering


developmental processes
Easily mapped within small genome
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Morphological patterns of growth are


markedly different between plants and
animals
Animal embryos become organized along
three axes and subdivide into segments
Form of higher plants has two key features

Root-shoot axis
Most growth occurs via cell division near the tips of
the shoots and the bottom of the roots
Growth occurs in a well-defined radial axis
e.g., Leaves and branches grow from the main
shoot
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Plant and animal development also differ


at the cellular level
Cell migration does not occur in plant
development
Plant development does not asymmetrically
deposited morphogens

An entirely new individual can be regenerated

from most types of somatic cells


Most plant cells are totipotent
Have the ability to produce an entire individual

Despite differences, the


underlying molecular
mechanisms of pattern
development still share
similarities with those of
animals

Development of seed plants

First post-fertilization cell division is


asymmetrical
Smaller apical cell
Gives rise to most of the embryo
Will later develop into the shoot of the plant
Larger basal cell
Will give rise to the root
Will give rise to extraembryonic tissue required for
seed formation

Development of seed plants

Basic organization of the plant has been


established by the heart stage
Approximately 100 cells

Development of seed plants

Shoot meristem will arise from a group of


cells located between the cotyledons
Precursors that will produce the plants shoot

Root meristem is located on the opposite


side
Will create the root

Meristems contain an organized group of


actively dividing stem cells
Stem cells retain their ability to divide and
differentiate into multiple cell types
Produce offshoots of proliferating cells as they
grow

e.g., Offshoots on shoot meristems are buds


Give rise to leaves and flowers

The shoot meristem is organized into


three areas
Organizing center
Central zone
Peripheral zone

The shoot meristem is organized into three


areas

Organizing center
Ensures the proper orientation of the meristem
Preserves the correct number of actively dividing stem

cells

Central zone
Undifferentiated stem cells are always maintained

Peripheral zone
Contains dividing cells that

will eventually differentiate


into plant structures
e.g., May form a bud, which
will produce a leaf or flower
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Organization of the Arabidopsis shoot


meristem

Controlled by two critical genes


WUS and CLV3

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Organization of the Arabidopsis shoot


meristem

WUS gene
Encodes a transcription factor expressed in the

organizing center
Induces adjacent cells in the central zone to become
undifferentiated stem cells
These stem cells then
turn on the CLV3 gene

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Organization of the Arabidopsis shoot


meristem

CLV3 gene
Activated by the WUS gene
Encodes a secreted protein
Binds to receptors in cells of the

peripheral zone
Prevents them from expressing the WUS gene
Limits area of WUS expression
Maintains a small population of stem cells at the
growing tip
Allows peripheral cells to differentiate and produce
specific structures
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Three main regions in the seedling

Apical region
Produces leaves

and flowers

Central region
Creates the stem

Basal region
Produces the roots

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Three main regions in the seedling


Apical, central, and basal regions
Each region develops differently

Unique cell division

patterns and distinct


morphologies

Each region expresses


different sets of genes

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Apical-basal-patterning genes

Category of genes important in


early stages of development
Defects cause dramatic effects in
one of the three main regions
e.g., A defective gurke gene results

in an embryo lacking apical


structures

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The first known homeotic


genes were described in
plants
Double flowers with stamens
replaced by petals were noted in
ancient Greece and Rome
Many homeotic mutations
affecting flower development
have been identified in
Arabidopsis
and the
snapdragon

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A normal Arabidopsis flower is composed


of four concentric whorls of structures
Outer whorl contains sepals
Second whorl is composed
of petals
Third whirl contains stamens

Produce male gametophyte

Innermost whorl contains


carpels
Produce female gametophyte

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Homeotic mutants of Arabidopsis have


undergone transformations of particular
whorls

e.g., Sepals transformed into carpels,


petals transformed into stamens, etc.

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ABC model for flower development

Four classes of genes govern the formation of


sepals, petals, stamens, and carpels
A, B, C, and Sepallata (SEP genes)

Flower structures are modified


leaves
The actions of various combinations

of these genes promotes the formation


of the various flower structures
Defective versions of A, B, and C
result in a flower composed solely
of leaves
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ABC model for flower development

Different combinations of genes are expressed


in each whorl
Leads to the formation of specific structures
Alterations in expression lead to the formation of

structures in aberrant locations

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ABC model for flower development

Genes A and C repress each others


expression
When one is lost, the other replaces it

Gene B functions independently

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ABC model for flower development

Loss of A
C is expressed in all whorls
Carpel-stamen-stamen-carpel arrangement

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ABC model for flower development

Loss of B
Petals and stamens cannot be made
Sepal-sepal-carpel-carpel arrangement

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ABC model for flower development

Loss of C
A is expressed in all four whorls
Sepal-petal-petal-sepal arrangement

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ABC model for flower development

Loss of SEP
Flower consists solely of sepals

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ABC model for flower development

Loss of A, B, and C
Flower consists solely of leaves

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ABC flower development in Arabidopsis

Two types of gene A


apetala1 and apetala2

Two types of gene B


apetala3 and pistillata

One type of gene C


agamous

Three SEP genes


SEP1, SEP2, and SEP3

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All of these plant homeotic genes encode


transcription factors
Proteins possess a DNA-binding domain and a
dimerization domain
Arabidopsis homeotic genes lack a sequence
similar to the homeobox found in animal
homeotic genes
Most of them (except apetala2) possess a
MADS box

Analogous to the homeobox


Promotes binding to specific DNA sequences
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Plant homeotic genes are part of a


hierarchy of gene regulation
Genes expressed within a flower bud
primordium produce proteins that activate the
expression of these homeotic genes
Products of homeotic genes
regulate the expression of other
genes
The products of these other genes
promote the formation of specific
flower structures

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