Embryo Development and

Chloroplast Biogenesis
 Plastids
• Plastids are a characteristic component of
plant cells
• Plastids are classified and named based
on the kinds of pigments they contain
• Each plastid is surrounded by two
membranes and internally the plastid has
a system of membranes which form
flattened sacs called thylakoids and a
ground (fluid) substance called stroma
Proplastids are small, colorless or
pale green undifferentiated plastids
that occur in egg cells,
meristematic cells of roots and
shoots - they will eventually develop
into other, differentiated plastids
such as the chloroplasts,
chromoplasts or leucoplasts
Chloroplasts are part of a large family of plastids
The life cycle of a flowering plant
 Light micrograph of the Electron micrograph
egg cell (EC) in showing a longitudinal
Median longitudinal section of the longitudinal section section of the egg cell (EC).
ovule and funicle, showing the
mature embryo sac and
constituent cells
 Endosperm
• The triploid cell divides repeatedly to form a
nutritive tissue called endosperm. This tissue
(like the yolk in animal eggs) stores nutrients
inside the seed for embryonic development,
seed germination, and early seedling growth.
 Embryogenesis
• In flowering plants, embryogenesis takes
place inside the ovule as the seed matures.

• Embryogenesis produces a tiny, simplified

plant.
 What Happens during Embryogenesis
• After fertilization, the zygote divides asymmetrically,
producing a large basal cell and a small apical cell.

• The basal (bottom) cell gives rise to the suspensor,

which anchors the embryo as it develops.

• The apical (top) cell gives rise to the mature embryo.

• The asymmetries in the basal and apical cells help

establish the apical-basal axis (top and bottom) of the
plant.
 What Happens during Embryogenesis
• Groups of cells called the shoot apical meristem (SAM) and root
apical meristem (RAM) form next.

• A meristem consists of undifferentiated cells that divide repeatedly,

with some daughter cells becoming specialized cells.

– Meristematic tissues produce cells in this way throughout the plant’s

life.

• Unlike animals, plant growth and development take place without

cell migration.

• Plant embryonic structures take shape because cell divisions occur

in precise orientations; the resulting cells exhibit differential
growth.
The zygote between 2 and 4 HAF (longitudinal sections).
Light and electron micrographs of the single cell and two-terminal cell embryos 12 and 18
HAF (longitudinal sections)
The heart stage embryo 66-84 HAF
Several steps are required for chloroplast biogenesis and
development

Non-randomly distributed chloroplast-containing cells are

seen as early as the globular stage of embryogenesis in
Arabidopsis – protodermal cells

In the heart stage of embryogenesis, chloroplast

containing cells are detected in epidermal cells as well as
a central region of the heart stage embryo, forming a
triangular septum of chloroplast-containing cells that
divides the embryo into three equal sectors.

Torpedo stage embryos have chloroplast-containing

epidermal cells and a central band of chloroplast-
containing cells in the cortex layer, just below the shoot
apical meristem.

In the walking-stick stage of embryogenesis,

chloroplasts are present in the epidermal, cortex and
endodermal cells.
 Interplay between two genomes

PLASTID
GENOME
~100 genes

Nucleus Import

TIC
PhANGs TOC
~3000 proteins
Chloroplast

cTP

30
 Interplay between two plastid RNA polymerases

PLASTID
GENOME

Transcription
NEP Plastid Encoded Polymerase

NEP Housekeeper genes

Nucleus Import

TIC
PhANGs TOC

Proplastid
NEP

cTP

31
 Interplay between two plastid RNA polymerases

PLASTID
GENOME
PEP Photosynthesis related genes

Transcription
NEP Plastid Encoded Polymerase

NEP Housekeeper genes

Nucleus Import

TIC
PhANGs TOC

Chloroplast
NEP

cTP

32
 Expression of plastid genes and plastid protein homeostasis

Translation
Nuclear-encoded
30S
PLASTID
GENOME 50S
Plastid-encoded
Assembly

Folding
Transcription

RNA processing Degradation

- Stability
- Editing
Import
- Splicing
- Co-translation
TIC
PhANGs TOC

NEP

cTP

33
 Expression of plastid genes and plastid protein homeostasis

Nuclear-encoded
30S
PLASTID
GENOME 50S
Plastid-encoded

Retrograde
signaling
TIC
PhANGs TOC

NEP

cTP

34
Fig. 2 Comprehensive dataset of 510 EMB genes of Arabidopsis. (a) Contributions of
research groups to the analysis of embryo-defective mutants included in the dataset. (b)
Confirmation of EMB gene identities. Genes were confirmed through molecular
complementation (MC), the analysis of multiple alleles (MA) in the absence of
molecular complementation, or some other experimental approach. Most of the genes
lacking confirmation are represented by Meinke laboratory mutants (MLM) rather than
community mutants (CM). Excludes 52 genes (Supporting Information Dataset S8E,F)
originally designated as EMB but later removed from the main dataset because of
uncertain gene identities or mutant phenotypes. (c) Terminal embryo phenotypes of
strong mutant alleles.
Fig. 3 Distribution of protein functions in theEMBdataset of Arabidopsis relative to
terminal embryo phenotype. Protein classes: 1, DNAsynthesis and repair; 2, RNA
synthesis and modification; 3, Protein synthesis; 4, Protein modification and transport;
5, Protein degradation; 6, Chromosome dynamics; 7, Transcriptional regulation; 8,
Signaling and regulatory proteins; 9, Energy and electron transport; 10, Metabolism; 11,
Cell structure, membrane function, and vesicle trafficking; 12, Other; 13, Uncertain and
unknown.
Is photosynthesis important for proper embryo development?
Collection of mutants with altered plastid ribosomes

PRPL 1
PRPL 4
Large
PRPL11
PRPL 27
PRPL 35
PRPL 24
PRPL 28

PRPS 1
Small PRPS 17
PRPS 20
PRPS21
The plastid ribosomal protein PRPS20, L1, L4, L27 and L35 are
essential for embryo development. PRPL28 is involved in greening of
embryos

51
 The thylakoid electron transport chain

Stroma

FNR
4 fotoni
4 fotoni 2H+ + 2NADP
6 H+ 2 NADPH
PQ Fd

CYT B6f

PSII O2 + 4H+
PSI
PC 4 e-
2H2O
Lume
 The thylakoid electron transport chain

Stroma

FNR
4 fotoni
4 fotoni 2H+ + 2NADP
6 H+ 2 NADPH
PQ Fd

CYT B6f

PSII O2 + 4H+
PSI
PC 4 e-
2H2O Lume
The thylakoidal complexes
The thylakoidal complexes
The chloro-embryos contain chlorophyll a and b

Chl a/b ratio is lower than in leaves and it is similar to shade plants adapted to low light and
green-enriched light

Sesbania sesban
PSII efficiency under different light intensities in chloro-embryos ( ) and leaves ( )

Larger antenna size in embryos than leaves and higher

ratio of PSI to PSII activity in developing chloroembryos
The thylakoidal complexes
Proper embryogenesis?
The cyclic electron transport
The cyclic electron transport: NDH and PGRL1-PGR5 complexes

da Shikanai 2007 Annu. Rev. Plant Biol.

Cold delays the establishment of a fully functional photosynthetic apparatus in cpna2-4 mutants
Photosynthetic apparatus functions properly in cpna2-4 seeds developing at 22 C whereas
exposure to cold decreases photosynthetic activity
Embryonic photosynthesis does not contribute to seed development
Perturbation of Embryonic photosynthesis leads to profound developmental defects starting
from early seedling development
Embryonic seed photosynthesis is important for
germination and the vigour of seedlings and mature plants