620 Part VI Renewal

Plastid number per cell ranges from more than 700 in

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
Figure 16.42 Double fertilization in maize. Two sperm are and survival of adventitious embryos is dependent on the
released from the pollen tube into a degenerated synergid. development of endosperm derived from normal double
One fuses with the egg and the second with the central cell. fertilization. This kind of apomixis is essentially a form
of somatic embryogenesis, equivalent to the
developmental pathway undertaken by cells in tissue
non-disjunction (‘not coming apart’), in which a pair of culture that have been induced to regenerate into plants.
chromosomes fails to separate correctly during meiosis I, Three families, the Asteraceae, Rosaceae and Poaceae,
meiosis II or mitosis. In lines of maize carrying B which collectively constitute only 10% of flowering plant
chromosomes, non-disjunction at the second pollen species, account for about 75% of gametophytic
mitosis results in one sperm cell receiving two or more B apomicts. Often, but not always, apomixis is associated
chromosomes while the other receives none. Sperm cells with self-incompatibility, perenniality and dehiscent
with B chromosomes fertilize egg cells more frequently fruits (i.e. those that split open at maturity).
than those without (in a ratio of up to 3:1). In both Gametophytic apomixis originates with a diploid
nuclear and cytoplasmic heterospermy, selectivity is embryo sac, which develops by mitosis of a diploid cell
believed to be the result of signaling between egg and with apomictic potential rather than from a haploid
sperm, but the molecular mechanisms remain to be megaspore (Figure 16.43). Endosperm development
discovered. may be either spontaneous (autonomous) or induced
Fertilization is the point at which organelle genomes by fertilization with a pollen sperm nucleus
are transmitted to the next generation, a phenomenon (pseudogamous).
called cytoplasmic inheritance. The predominant type of Two pathways of gametophytic apomixis, diplospory
cytoplasmic inheritance is uniparental maternal, in and apospory, are recognized, based on the cell type that
which the genotypes of plastids and mitochondria in the gives rise to the diploid embryo sac. In diplospory
progeny are derived exclusively from the female parent. (Figure 16.43B), the megaspore mother cell may initiate
Uniparental paternal inheritance of plastids is found in meiosis but abort it at an early stage and switch to mitotic
a significant minority of flowering plants. Inheritance of development of the embryo sac, a condition referred to
mitochondria via the paternal line has been described for as meiotic diplospory. Dandelion (Taraxacum) is an
alfalfa (Medicago sativa), Populus and Brassica napus. The example of such a diplosporous apomict. An alternative
plastids of the egg cell are usually of variable size and mechanism is mitotic diplospory, where the MMC
shape, contain some starch grains and lamella-like proceeds directly to form the unreduced embryo sac by
structures, and tend to cluster around the nucleus. 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 into major crop species has the prospect of
by members of the hawkweed genus (Hieracium). In revolutionizing crop breeding. Many of the most
apospory the embryo sac is derived from one or more productive and stress-resistant agricultural and
somatic ovule cells, called aposporous initials. Diploid horticultural crop varieties are hybrids. The superior
embryo sacs can differentiate from aposporous initials at performance of the progeny of a cross between two
various times during ovule development and may coexist inbred parents is called heterosis or hybrid vigor.
with meiotically haploid embryo sacs in the same ovule Conversely, the consequence of self-pollinating hybrids
or may continue to develop while the haploid sexual over several generations is inbreeding depression, that
embryo sac degenerates (Figure 16.43C). is, progressive reduction in heterozygosity and vigor. An
Apomixis has been called an evolutionary dead end or example of the exploitation of heterosis in a major crop
blind alley because of its postulated association with is maize. The discoverer of heterosis in maize was
genetic uniformity, low adaptive potential and the Charles Darwin, who noted that the progeny of
accumulation of deleterious mutations. On the other cross-pollinated maize were 25% taller than the progeny
hand apomixis, like other forms of asexual reproduction of inbred parents. Hybrid varieties were first developed
(discussed in detail in Section 11.4.2 and in Chapter 17), by maize breeders early in the 20th century. Since that
is an abundant and widely distributed attribute that time their use has increased, together with crop yields,
clearly confers adaptive and evolutionary advantages in until today hybrids are planted in about 95% of maize
dynamic natural populations. This makes the genetic acreage in the United States, and two-thirds worldwide.
regulation of apomixis a subject of ecological interest. Figure 16.44 shows that the heterotic effect is apparent
Apomixis has also attracted agronomic and not only in grain yield at maturity but also from the
biotechnological attention because introducing the trait earliest phase of seedling development.
622 Part VI Renewal

Inbred A

Inbred B
Inbred A Hybrid A × B Inbred B 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 segregation, meiosis is totally replaced by mitosis. Such
breed true, and suffer from inbreeding depression if so-called MiMe plants produce diploid male and female
propagated by conventional methods of seed gametes that are genetically identical to their parent, and
multiplication. It is therefore necessary to recreate the ploidy doubles at each generation. As more detailed
hybrid each season by crossing inbred parents, an molecular understanding of meiosis and apomixis is
expensive and time-consuming business. This is where acquired, new tools and approaches such as MiMe plants
interest in apomixis comes in, because it offers the are becoming available to help the biotechnologist and
prospect of creating hybrids that propagate asexually via plant breeder reach the goal of being able to manipulate
seeds and in which desirable traits are therefore the reproductive systems of crops at will.
transmitted faithfully to the progeny. The genus
Tripsacum (gamagrass), which is related to Zea, includes

16.6 Seed and fruit

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
development
cross with maize and there have been numerous
attempts to transfer the apomictic locus into maize by Pollination and double fertilization are followed by the
exploiting the interfertility between the two species. So development of major structures of the seed: the embryo,
far success has been limited, at least in part because of endosperm and seed coat. The seed coat originates in
low rates of recombination between the genomes of the ovule integuments, the maternal tissues that provide
two species, but backcross programs to integrate the protection and facilitate nutrient transfer to the
apomixis locus into the maize genetic background developing embryo they surround. A fruit (see
continue. Some authorities believe the organization of Figure 1.22) is defined as a structure derived from an
genes for apomixis is too complex for intergeneric ovary and bearing or containing seeds. It may be simple
transfer of the trait by conventional crossing ever to be (originating as a single ovary), aggregate (from several
possible. separate ovaries of a single flower) or multiple (derived
An alternative strategy is to intervene in the process of from an inflorescence). The winged fruit of Acer spp. is
meiosis in the MMC by genetic engineering. An example an example of a simple type. Raspberries (Rubus idaeus)
of this approach is a recent study that identified an are aggregate fruits. Hop (Humulus) is a multiple fruit.
Arabidopsis gene named OSD1 (OMISSION OF The structures of some fruits include tissues other than
SECOND DIVISION 1). The products of meiosis in osd1 those of the gynoecium, in which case they are called
mutants are dyad rather than tetrad spores. In triple false fruits or pseudocarps. Strawberry (Fragaria), in
mutants consisting of osd1 with a gene that eliminates which the red fleshy part is derived from the floral
recombination and another that modifies chromatid receptacle, is a false fruit (see Figure 6.5A).