Gene transfer, asexual reproduction, sexual reproduction, and apomixis Plant Reproduction  Gene transfer, asexual reproduction, sexual reproduction, and apomixis viachicago.wordpress.com birdsandbloomsblog.com tinyfarmblog.com

Asexual reproduction The clone is immortal Example: Allium sativum “As far as we know, garlic in cultivation throughout history has only been propagated asexually by way of vegetative cloves, bulbs, and bulbils (or topsets), not from seed. These asexually propagated, genetically distinct selections of garlic we cultivate are more generally called "clones". Yet this asexual lifestyle of cultivated garlic forgoes the possibility of combining traits proffered by interpollinating diverse parental stocks.” Source: http://www.ars.usda.gov/Research/docs.htm?docid=5232

Asexual reproduction The clone is immortal Example: Populus tremuloides The world's heaviest living thing 1 clone in the Wasatch Mountains of Utah 47,000 stems of genetically identical aspen trees Total weight: 6 million kilograms Aspen is dioecious species - this clone is one big male source: http://waynesword.palomar.edu/ww0601.htm#aspen

Sexual reproduction Advantages > disadvantages Genetic variation: Allele exchange via cross-pollination New combinations of alleles via meiosis Purge deleterious mutations Stay ahead in the host-pathogen “arms race” Potential adaptation to a changing climate

Sexual reproduction Advantages > disadvantages In a dioecious species, half the reproductive effort is wasted in producing males Meiosis produces some "unfit" combinations of alleles Cross-pollinated plants may be subject to environmental conditions unfavorable to pollination Finding a mate

Alternation of generations In plants there is an alternation of the gametophytic (n) and sporophytic (2n) generations (Gametophytic = haploid)

Alternation of generations The sporophytic generation may be diploid (2n = 2x) or polyploid (2n = _x) VAVA VAVAVBVB VAVAVBVBVDVD 2n = 2x = 14 30,000 genes 2n = 4x = 28 60,000 genes 2n = 6x = 42 90,000 genes 1 pair homologous chromosomes 0 sets of homoeologous chromosomes AA 2 pairs of homologous chromosomes 2 sets of homoeologous chromosomes AABB 3 pairs of homologous chromosomes 3 sets of homoeologous chromosomes AABBDD A A B B A A A A B B D D

Angiosperm reproductive organs and gamete formation Development of the female gametophyte Reproductive structure: Ovule(s), style, stigma http://www.extension.org/pages/32204/stigma

Angiosperm reproductive organs and gamete formation Development of the female gametophyte Megaspore mother cell (MMC)  MMC undergoes meiosis Of 4 megaspores produced 1 survives (most species) Three post-meiotic mitoses   1 2 3

Angiosperm reproductive organs and gamete formation Development of the female gametophyte The 8-nucleate embryo sac (1 egg, 2 synergids, 2 primary endosperm nuclei, 3 antipodals)   Source: yougems.reflectionsinfos.com lima.ohio-state.edu

Angiosperm reproductive organs and gamete formation Development of the female gametophyte Pollinator attraction: Petals, nectaries, etc. Source: arizonabeetlesbugsbirdsandmore.blogspot.com

Angiosperm reproductive organs and gamete formation Development of the male gametophyte Reproductive structures: Anthers; pollen within anthers

Angiosperm reproductive organs and gamete formation Development of the male gametophyte Pollen mother cell (PMC)    PMC undergoes meiosis Meiosis gives a tetrad of microspores Note, this is different than ♀ Meiosis 1 Meiosis 2

Angiosperm reproductive organs and gamete formation Development of the male gametophyte The first mitosis gives vegetative and generative nuclei; at the second mitotic division, the generative nucleus gives 2 sperms. mitosis mitosis mitosis mitosis mitosis mitosis mitosis mitosis

The pollen pathway and double fertilization The stigma is the site of pollen recognition Pollen germinates and the vegetative (tube nucleus) grows through the style to the ovule The two sperm use the tube as conduit

The pollen pathway and double fertilization Double fertilization: One sperm fertilizes the egg to give the 2n embryo, the other fertilizes the polar nuclei to give the 3n endosperm antipodals ♀ ♀ 3n endosperm polar nuclei ♀ ♀ ♂ ♀ ♂ ♀ egg 2n embryo synergids

The pollen pathway and double fertilization After double fertilization, there are at least four independent and genetically distinct generations coexisting in the seed: maternal sporophyte diploid tissue maternal gametophyte haploid tissue offspring sporophyte diploid tissue fusion of male (1) and female (2) gametophyte to form triploid tissue

A review… V v sporophytic generation v v V V V v v V V V V V V V V V MMC V v V V v v v V V v v v v v v v v v PMC V v v V V v sporophytic generation v v V V v V v V v V v v V V

A review of sexual reproduction in an F1 plant, heterozygous for the Vv locus + = v v v V V V V V V V v V v + = V V V V V V V V V v VV Vv vv + = V V V v v v v v v v V v V + = v v v v v v Note: At this point in the figure, the antipodals and synergids are deleted and only the fertilized endosperm nuclei (now 3n) and fertilized egg (now 2n) are shown. Only the fertilized egg is carried to the Punnett square.

Seeds without sex: Apomixis Apomixis involves parthenogenesis – development of an egg without fertilization, as opposed to parthenocarpy – development of fruits (seedless) without fertilization Implication - Apomixis allows for seed propagation of a heterozygote – genetically identical from generation to generation. Obligate:   Facultative: 

Seeds without sex: Apomixis Hand and Koltunow. 2014. Genetics 197: 441-450.

Seeds without sex: Apomixis Economic implications of apomixis Why would apomixis be, or not be, of commercial interest to purveyors of hybrid seeds?  If transgenic apomicts are developed and released, and if apomixis is dominant, it will be necessary to control pollen flow – if not, there will be frozen gene pools Evolutionary implications of apomixis Obligate vs. facultative

Seeds without sex: Apomixis Prevalence 400 species ; 40 families; Common in Poaceae, Asteraceae, Rosaceae Examples Tripsacum Poa pratensis Pennisetum Dandelion (Taraxacum spp) Rubus

Seeds without sex: Apomixis How it works no (or altered) meiosis to produce unreduced female gametophyte (embryo sac) no fertilization – but do get autonomous embryo formation may have autonomous endosperm development, or endosperm development may be triggered by fertilization. Most cases it is triggered by fertilization (pseudogamy = fertilization of central cell)  

Seeds without sex: Apomixis Genetic basis Apomixis loci and candidate genes Lots of breeding effort; little success Epigenetically regulated?

The Rubus armeniacus case study Himalayan (Armenian) blackberry

Is the Himalayan blackberry the perfect weed? Designing the perfect invasive Attribute Description Flower Hermaphroditic Pollination biology Self and outcross Apomixis Facultative Seeds Small and numerous Vegetative propagation Yes Ploidy level Polyploid Protection Thorns Attraction Tasty fruit Was Luther Burbank the “father” of this perfect weed? promo.idahopotato.com