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Development of

Flower
Flower: report outline

◉ Stages of Development
• Structural Organization of the Inflorescence Meristem and Origin
of the Flower Meristem
• Floral Organ Primordia
• The stages of floral development
◉ The ABC Model
• What is the ABC Model?
• What are the ABC genes and how do they work?
• What do the ABC genes regulate?
• What about roses, tulips, and daisies?
• What about Pine trees and non-flowering plants?

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The basic floral architecture is


mostly conserved among the so-
called core eudicots, that makes
up over 73% of extant flowering
plants
(Drinnan et al., 1994)


Stages of Flower
1 Development

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Arabidopsis
Flower Model

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Structural Organization of the
Inflorescence Meristem and Origin
of the Flower Meristem

During the vegetative phase of the Arabidopsis life cycle, the shoot apical
meristem (SAM) produces leaves on its flanks and on transition to flowering, the
shoot bolts and the SAM becomes the inflorescence shoot apical meristem (IM).
On bolting, some of the pre-existing leaf primordia become cauline leaves
subtending lateral inflorescence shoots (paraclades) and the shoot apex starts to
produce flowers. A primary IM produces lateral meristems that may go on to
produce flowers or secondary inflorescences.
Arabidopsis inflorescences are subtended by fully developed bracts, but
flowers only by rudimentary ones. It is generally said that the IM generates the floral.

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Structural Organization of the
Inflorescence Meristem and
Origin of the Flower Meristem

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Floral Organ Primordia

Once a flower primordium is initiated, the geometry changes and a rapid


and coordinated burst of cell expansion and division occurs in three dimensions
generating a concentric group of cells as an almost spherical flower primordium,
from which all floral tissues are derived.
The regulation of cell divisions during early and late stages of flower
development seems to depend upon different mechanisms. Early in flower
development, when the floral meristem of Arabidopsis is divided into four
concentric rings (each with a characteristic multigenic expression profile), cell
division patterns depend upon the cell's radial position in the floral meristem, and
not on the future identity of the floral organ to be formed in each ring.

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Floral Organ Primordia

After stage 6, during organogenesis, the ABC homeotic genes seem to


control the rate and orientation of cell divisions. As a result, the continuity of the
concentric rings is broken giving distinct floral organ primordia within each
whorl, then cells sub differentiate into distinct types within each organ. The
initiation and identity of floral organs are also regulated by different and largely
independent molecular modules. This is suggested, for example, by the fact that
conversion of petals into sepal-like organs in mutant plants does not alter the
number of cells involved in their initiation.
Tissues of floral organs are organized according to coordinated patterns
and rates of cell division in the different cell layers of the meristem that
dynamically acquire distinct fates.
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The stages of floral
development

Stage 1 Stage 2
The first sign of flower primordium During this stage, the hemispherical primordium
formation is the bulging of the peripheral continues to grow forming almost a right angle
surface of the IM in a lateral direction. It is with the surface of the SAM, which itself
hypothesized that a lateral protrusion lengthens and widens rebuilding the portion of
formed during bulging is a rudimentary the periphery that has been used for
bract. primordium formation (Figures 3 and 4A–B).
During early stages of flower development At this stage the flower primordium becomes
in Arabidopsis, two types of primordia clearly delimited from the IM, and starts to grow
(bract and flower primordium proper) and larger very quickly in all directions (Figures 3
organ boundaries are observed. and 4A–B).
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Stage 1 and 2

Figure 2A Figures 4A–B


Stage 1 Stage 2

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The stages of floral
development
Stage 3 Stage 4
This stage begins when sepal primordia become During this stage, the elongation
visible. By now the flower primordium is 30–35
of the pedicel continues
µm in diameter and is becoming stalked with an
incipient pedicel. It has also started to grow concurrently with an increase in
vertically. the diameter of the developing
The two lateral (I) sepal primordia appear first, but flower primordium to 65–70 µm.
are soon outgrown by the abaxial (ab) then the The medial sepal primordia have
adaxial (ad) sepal primordia. Sepal primordia arise already partly overtopped the
initially as ridges that lengthen and curve inwards remaining floral meristem.
until they begin to overtop the remaining dome-
shaped portion of the flower primordium (Figures
3 and 4C–D).
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Stage 3 and 4

Figures 4C–D Figure 4E


Stage 3 Stage 4

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The stages of floral
development
Stage 5 Stage 6
This stage is when the petal and stamen The sepals grow to completely cover the
primordia become visible. Primordia of floral bud and the primordia of the four long
the four medial (long) stamens are first stamens bulge out and become distinct
seen as wide outgrowths on the flanks of from the central dome of cells that comprise
the central dome of the FM. the FM. The two lateral stamen primordia
The four petal primordia that arise arise slightly lower on the dome and
between the sepals close to their base are develop later. The petal primordia grow
just visible during this stage. The two somewhat but are still relatively small. A rim
lateral (short) stamens develop from around the central dome of the flower
primordia that appear later during this primordium now begins to grow upward to
stage (Figures 3 and 4F–G). produce an oval tube that will become the
gynoecium (Figures 3 and 4H–I).
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Stage 5 and 6

Figures 4F–G Figures 4H–I


Stage 5 Stage 6

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The stages of floral
development
Stage 7 Stage 8
This stage begins when the growing The beginning of stage 8 is defined by
primordia of the long stamens become another landmark in stamen
development: anther locules are visible
stalked at their base. The stalks give rise
as convex protrusions on the inner
to the filaments, and the wider upper (adaxial) surface of the long stamens.
region to the anthers.
At this stage stamens are 55–60 µm long
By this stage, petal primordia have most of which is the developing anther.
become hemispherical although they are Locules also appear soon after in the
still relatively small (ca. 25 µm in short stamens. Petal growth now
diameter; Figures 3 and 5A–B). accelerates and petal primordia become
apparent (Figures 3 and 5C–E).
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Stage 7 and 8

Figures 5A–B Figures 5C–E


Stage 7 Stage 8

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The stages of floral
development
Stage 9 Stage 10
This stage begins when the petal primordia
The rapidly growing petals reach
elongate. There is a rapid lengthening of all
organs especially of petals that acquire a
the top of the lateral stamens. The
tongue-like shape and increase in length from cap of papillae that will constitute
about 45 µm to up to 200 µm. Nectary glands the stigma starts to form at the
appear and the stamens grow rapidly. By the top of the gynoecium (Figures 3
end of stage 9, the medial stamens are around and 5H–I).
300 µm long. Most of this growth occurs in
the anther region, which still accounts for over
80% of their total length. At this stage the
floral bud remains completely closed (Figures
3 and 5F–G). 18
Stage 9 and 10

Figure 5F–G Figure 5H–I


Stage 9 Stage 10

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The stages of floral
development
Stage 11 Stage 12
This stage begins when the upper Petals continue to lengthen relatively
surface of the gynoecium develops rapidly. Lateral sepals continue to grow
while the stamens and gynoecium lengthen
stigmatic papillae (Figures 3 and
coordinately. The anthers have almost
6A–C) although their outward reached their mature length of 350–400 µm
growth is limited at first to regions and the filaments now lengthen rapidly. The
not in contact with the overlapping upper part of the gynoecium differentiates
sepals. By the end of this stage petal into the style (Figure 6D) and a sharp
primordia reach the top of the boundary separates it from the cap of
medial stamens. stigmatic papillae. Stage 12 ends when the
sepals open (Figures 3 and 6D–F).
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Stage 11 and 12

Figure 6A-C Figure 6D-F


Stage 11 Stage 12

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The stages of floral
development
Stage 13 Stage 14
Petals become visible between the sepals and This is also defined as the stage zero hours
continue to elongate rapidly. The stigma is after flowering (0 HAF), and it marks the
receptive at this stage (Figures 3 and 6G–H). beginning of silique (the fertilized pistil or
Stamen filaments extend even faster so the fruit) and seed development. Cells in the
stamens outstrip the gynoecium in length and exocarp continue to divide anticlinally and
self-pollination takes place. The gynoecium is expand longitudinally in the replum and
now mature and its three distinct regions can the valve, where there is also some
be distinguished: an apical stigma, a style, and expansion in other directions. There is also
a basal ovary. After pollination, pollen tubes division and expansion in the mesocarp
grow to fertilize the ovules, the stamens and many chloroplasts develop (Figures 3
extend above the stigma, and furrows at both and and6I6I).
valve/replum boundaries appear. 22
Stage 13 and 14

Figure 6G-H Figure 6I


Stage 13 Stage 14

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The stages of floral
development

Stage 15 Stage 16
The stigma extends above the long At this stage the silique is twice as
anthers. In the carpel walls, cell long as a stage-13 pistil. Petals and
division and expansion continue. sepals wither and tissues in the
The medial vascular bundles silique continue expanding (Figures 3
continue to grow and xylem and and6L6L).
lignifies, while the lateral bundles
branch out through the mesocarp
(Figures 3 and 6J–K).

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Stage 15 and 16

Figure 6J-K Figure 6L


Stage 15 Stage 16

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The stages of floral
development
Stage 17 Stage 18
The silique begins to yellow from the tip to
This stage is defined by the abscission the base. One of the endocarp cell layers
of the senescent floral organs from the (the second from the inside) lignifies
silique, ∼2 days after fertilization. The further, and the inner endocarp cell layer
green silique grows to reach its final disintegrates, while the mesocarp begins to
length and matures, a phase lasting dry out. It has been suggested that
about 8 days making this the longest lignification may contribute to the silique
stage. The dehiscence zone also shattering process, acting in a springlike
differentiates (Figures 3, 7A and 7E) manner to create mechanical tensions
(Figures 3 and 7B).

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Stage 17 and 18

Figure 7B
Stage 18

Figure 7A and 7E
Stage 17

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The stages of floral
development

Stage 19 Stage 20
The valves begin to separate At this stage the valves become
from the dry silique, separated from the dry silique
apparently owing to the lack and the mature seeds are ready
of cell cohesion at the to be dispersed (Figures 3, 7D
separation layer.(Figures 3 and and 7F).
and7C7C).

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Stage 19 and 20

Figure 7C
Stage 19

Figure 7D and 7F
Stage 20

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Figure 3 30
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2 The ABC Model

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What is the ABC Model?

It is a model showing the homeotic mutations


affecting floral organ identity had been known
for centuries, it was the systematic analyses of
these mutations, and of the phenotypes
produced by double and triple mutants, that
proved to be critical in developing the ABC
model.

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What is the ABC Model?

Wild type Arabidopsis flower (A), color coded in (B) to demarcate the sepals (red), petals
(purple), stamens (green) and carpels (yellow). (C) A cartoon version of an Arabidopsis flower,
with the domains of ABC gene function shown below. The function of A alone specifies sepal
identity (red) in the first whorl, while a combination of A (red) + B (blue) function specifies petal
identity (purple) in the second whorl. The combination of B (blue) + C (yellow) function specifies
stamen identity (green) in the third whorl, while C function alone specifies carpels (yellow) in the
fourth whorl. (D–F) How the ABC model explains different mutant phenotypes.
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What is the ABC Model?

(D) The elimination of A function results in flowers composed of carpels in the first whorl,
stamens in the second and third whorls, and carpels in the fourth whorl; the lack of A function
in the first two whorls results in ectopic expression of C function in those regions. (E) A flower
lacking B function consists of sepals in the first and second whorls and carpels in the third and
fourth whorls. (F) A flower lacking C function consists of sepals in the first whorl, petals in the
second and third whorls, and sepals in the fourth whorl. 35
What are the ABC genes and
what do they regulate?

The ABC genes


◉ ABC genes are class of homeotic
genes
◉ ABC genes works together in
flower development

KEY NOTE:
◉ A and C genes are equally
dominant
◉ B genes always associates with A
and C genes
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What are the ABC genes and
what do they regulate?

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What do the ABC Genes
Regulate?

The ABC Model of flower development in angiosperm


demonstrates the presence of the three classes of genes
that regulate the development of floral organs. Class A, B
and C genes are homeotic genes. They induce the organs to
develop in their respective whorls.

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What do the ABC Genes
Regulate?
◉ The ABC proteins are responsible for
turning on and turning off the
expression of different panoplies of
genes.
◉ Little is known of the genes and
processes regulated by the ABC gene
products.
◉ B proteins control cell division genes
that act to shape the petal; that petal
epidermal cells are sculpted in part
by genes that are turned in in
response to B gene activity. 41
What about roses, tulips and
daisies?

Despite these morphological variations, ABC genes have been identified and
shown to function within each tiny daisy flower in a very similar manner to
their Arabidopsis counterparts
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What about roses, tulips and
daisies?

Cultivated roses have many extra whorls of petals while


tulips lack obvious sepals
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What about roses, tulips and
daisies?

Both of these flowers


can be explained by
simple shifts in the
patterns of
expression of ABC
genes.

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Non-flowering plants
Gymnosperm

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Non-flowering plants

ABC Genes

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Non-flowering plants

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Non-flowering plants

Vascular and Non-vascular plants

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Thanks!
Any questions ?

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