620 Part VI Renewal
Plastid number per cell ranges from more than 700 in
Figure 16.42 Double fertilization in maize. Two sperm are
released from the pollen tube into a degenerated synergid.
One fuses with the egg and the second with the central cell.
non-disjunction (‘not coming apart’), in which a pair of
chromosomes fails to separate correctly during meiosis I,
meiosis II or mitosis. In lines of maize carrying B
chromosomes, non-disjunction at the second pollen
mitosis results in one sperm cell receiving two or more B
chromosomes while the other receives none. Sperm cells
with B chromosomes fertilize egg cells more frequently
than those without (in a ratio of up to 3:1). In both
nuclear and cytoplasmic heterospermy, selectivity is
believed to be the result of signaling between egg and
sperm, but the molecular mechanisms remain to be
discovered.
Fertilization is the point at which organelle genomes
are transmitted to the next generation, a phenomenon
called cytoplasmic inheritance. The predominant type of
cytoplasmic inheritance is uniparental maternal, in
which the genotypes of plastids and mitochondria in the
progeny are derived exclusively from the female parent.
Uniparental paternal inheritance of plastids is found in
a significant minority of flowering plants. Inheritance of
mitochondria via the paternal line has been described for
alfalfa (Medicago sativa), Populus and Brassica napus. The
plastids of the egg cell are usually of variable size and
shape, contain some starch grains and lamella-like
structures, and tend to cluster around the nucleus.
Plumbago to as few as eight in Daucus (carrot) species.
16.5.8 Apomixis, asexual
reproduction through seeds,
occurs in a large number of
taxa and is a target trait for
crop breeding
In some plant species, seeds can be produced asexually as
well as by sexual reproduction. The processes by which
seeds are produced in the absence of fertilization of an
egg cell by a sperm are called apomixis (also termed
agamospermy); a plant that reproduces by apomixis is
called an apomict. Apomixis pathways are classified as
gametophytic or sporophytic according to the source of
the cells that develop into an embryo. The apomictic
habit has evolved independently several times and occurs
in more than 400 taxa. Sporophytic apomixis, also called
adventitious embryony, is so-named because embryos
arise spontaneously from cells of ovule integuments or
from nucellar cells that are adjacent to a haploid embryo
sac. It is common among citrus species. The maturation
and survival of adventitious embryos is dependent on the
development of endosperm derived from normal double
fertilization. This kind of apomixis is essentially a form
of somatic embryogenesis, equivalent to the
developmental pathway undertaken by cells in tissue
culture that have been induced to regenerate into plants.
Three families, the Asteraceae, Rosaceae and Poaceae,
which collectively constitute only 10% of flowering plant
species, account for about 75% of gametophytic
apomicts. Often, but not always, apomixis is associated
with self-incompatibility, perenniality and dehiscent
fruits (i.e. those that split open at maturity).
Gametophytic apomixis originates with a diploid
embryo sac, which develops by mitosis of a diploid cell
with apomictic potential rather than from a haploid
megaspore (Figure 16.43). Endosperm development
may be either spontaneous (autonomous) or induced
by fertilization with a pollen sperm nucleus
(pseudogamous).
Two pathways of gametophytic apomixis, diplospory
and apospory, are recognized, based on the cell type that
gives rise to the diploid embryo sac. In diplospory
(Figure 16.43B), the megaspore mother cell may initiate
meiosis but abort it at an early stage and switch to mitotic
development of the embryo sac, a condition referred to
as meiotic diplospory. Dandelion (Taraxacum) is an
example of such a diplosporous apomict. An alternative
mechanism is mitotic diplospory, where the MMC
proceeds directly to form the unreduced embryo sac by
mitosis without an initial meiotic phase. The second
 Chapter 16 Flowering and sexual reproduction 621

Figure 16.43 Mechanisms of gametophytic apomixis compared with normal sexual fertilization. In each case, female
reproduction occurs in the nucellus of the ovule, where a single cell typically becomes the megaspore mother cell (MMC). (A) Sexual
fertilization. Meiosis of the 2n MMC gives rise to the four haploid cells of the reduced embryo sac. Double fertilization results in
diploid embryo and triploid endosperm. (B) Diplosporous apomixis. In this type, there is no meiotic reduction of the MMC. The
embryo is 2n without fertilization and only the sperm cell nucleus destined to form endosperm penetrates the embryo sac. (C) In
apospory, the MMC undergoes meiotic reduction, but one or more embryo sacs form from 2n aposporous initials adjacent to the
MMC or megaspore. Endosperm may result from the fusion of one sperm nucleus with the polar nucleus. In apospory and
diplospory, endosperm may also form in the absence of a pollen nucleus.

form of gametophytic apomixis is apospory, exemplified
by members of the hawkweed genus (Hieracium). In
apospory the embryo sac is derived from one or more
somatic ovule cells, called aposporous initials. Diploid
embryo sacs can differentiate from aposporous initials at
various times during ovule development and may coexist
with meiotically haploid embryo sacs in the same ovule
or may continue to develop while the haploid sexual
embryo sac degenerates (Figure 16.43C).
Apomixis has been called an evolutionary dead end or
blind alley because of its postulated association with
genetic uniformity, low adaptive potential and the
accumulation of deleterious mutations. On the other
hand apomixis, like other forms of asexual reproduction
(discussed in detail in Section 11.4.2 and in Chapter 17),
is an abundant and widely distributed attribute that
clearly confers adaptive and evolutionary advantages in
dynamic natural populations. This makes the genetic
regulation of apomixis a subject of ecological interest.
Apomixis has also attracted agronomic and
biotechnological attention because introducing the trait
into major crop species has the prospect of
revolutionizing crop breeding. Many of the most
productive and stress-resistant agricultural and
horticultural crop varieties are hybrids. The superior
performance of the progeny of a cross between two
inbred parents is called heterosis or hybrid vigor.
Conversely, the consequence of self-pollinating hybrids
over several generations is inbreeding depression, that
is, progressive reduction in heterozygosity and vigor. An
example of the exploitation of heterosis in a major crop
is maize. The discoverer of heterosis in maize was
Charles Darwin, who noted that the progeny of
cross-pollinated maize were 25% taller than the progeny
of inbred parents. Hybrid varieties were first developed
by maize breeders early in the 20th century. Since that
time their use has increased, together with crop yields,
until today hybrids are planted in about 95% of maize
acreage in the United States, and two-thirds worldwide.
Figure 16.44 shows that the heterotic effect is apparent
not only in grain yield at maturity but also from the
earliest phase of seedling development.
622 Part VI Renewal
Inbred A Hybrid A × B
A B
Figure 16.44 Heterosis in maize. (A) Grain yield in the hybrid between A and B exceeds that of either inbred parent. (B) Hybrid
vigor is expressed from the earliest stages of seedling growth.
The downside to F1 hybrid varieties is that they do not
breed true, and suffer from inbreeding depression if
propagated by conventional methods of seed
multiplication. It is therefore necessary to recreate the
hybrid each season by crossing inbred parents, an
expensive and time-consuming business. This is where
interest in apomixis comes in, because it offers the
prospect of creating hybrids that propagate asexually via
seeds and in which desirable traits are therefore
transmitted faithfully to the progeny. The genus
Tripsacum (gamagrass), which is related to Zea, includes
apomictic species. Tripsacum dactyloides diploids are
sexual, but tetraploid genotypes are pseudogamous
diplosporous apomicts. Diplospory in Tripsacum is
inherited as a single dominant locus. T. dactyloides will
cross with maize and there have been numerous
attempts to transfer the apomictic locus into maize by
exploiting the interfertility between the two species. So
far success has been limited, at least in part because of
low rates of recombination between the genomes of the
two species, but backcross programs to integrate the
apomixis locus into the maize genetic background
continue. Some authorities believe the organization of
genes for apomixis is too complex for intergeneric
transfer of the trait by conventional crossing ever to be
possible.
An alternative strategy is to intervene in the process of
meiosis in the MMC by genetic engineering. An example
of this approach is a recent study that identified an
Arabidopsis gene named OSD1 (OMISSION OF
SECOND DIVISION 1). The products of meiosis in osd1
mutants are dyad rather than tetrad spores. In triple
mutants consisting of osd1 with a gene that eliminates
recombination and another that modifies chromatid
Inbred A
Inbred B
Inbred B Hybrid A × B
segregation, meiosis is totally replaced by mitosis. Such
so-called MiMe plants produce diploid male and female
gametes that are genetically identical to their parent, and
ploidy doubles at each generation. As more detailed
molecular understanding of meiosis and apomixis is
acquired, new tools and approaches such as MiMe plants
are becoming available to help the biotechnologist and
plant breeder reach the goal of being able to manipulate
the reproductive systems of crops at will.
16.6 Seed and fruit
development
Pollination and double fertilization are followed by the
development of major structures of the seed: the embryo,
endosperm and seed coat. The seed coat originates in
ovule integuments, the maternal tissues that provide
protection and facilitate nutrient transfer to the
developing embryo they surround. A fruit (see
Figure 1.22) is defined as a structure derived from an
ovary and bearing or containing seeds. It may be simple
(originating as a single ovary), aggregate (from several
separate ovaries of a single flower) or multiple (derived
from an inflorescence). The winged fruit of Acer spp. is
an example of a simple type. Raspberries (Rubus idaeus)
are aggregate fruits. Hop (Humulus) is a multiple fruit.
The structures of some fruits include tissues other than
those of the gynoecium, in which case they are called
false fruits or pseudocarps. Strawberry (Fragaria), in
which the red fleshy part is derived from the floral
receptacle, is a false fruit (see Figure 6.5A).