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Plant Breeding Systems: evolution and diversity of reproduction modes in flowering plants Patrik MRÁZ: Special lesson within of 'Introduction to the study of biodiversity' course

Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

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Page 1: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Plant Breeding Systems:

evolution and diversity of reproduction modes in flowering plants

Patrik MRÁZ: Special lesson within of 'Introduction to the study of biodiversity' course

Page 2: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mating systems in plants and animals: a comparison

Introduction Templeton et al, BiolLett 2012

● absence of active mate-choice (no central nervous system

Page 3: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mating systems in plants and animals: a comparison

Introduction Templeton et al, BiolLett 2012

● absence of active mate-choice (no central nervous system)

Page 4: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mating systems in plants and animals: a comparison

Introduction http://www8.nos.noaa.gov/estuaries101/About/LearnMore.aspx?ID=356

● asexual reproduction more frequent (totipotency: genet and ramets)

Page 5: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mating systems in plants and animals: a comparison

Introduction http://bioweb.uwlax.edu/bio203/f2011/proft_paxt/reproduction.htm

● alternation of gametophytic and sporophytic generation

Page 6: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mating systems in plants and animals: a comparison

Introduction

● alternation of gametophytic and sporophytic generation

harmful mutations expressed in the gametophyte are not trasfered to sporophytic generation: more effective selection against the genetic load > selfing quite common

Page 7: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mating systems in plants and animals: a comparison

Introduction

● predominance of hermaphrodites (72% in flowering plants)

Page 8: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Introduction

● hermaphrodity versus unisexuality

● self-pollination versus cross-pollination

● self-fertilization versus cross-fertilization

● sexuality versus asexuality

Mating systems in plants and animals: a comparison

Plants: variable reproductive strategies

Page 9: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Reproductive strategies options

Introduction

sexual asexual

Page 10: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Reproductive strategies options

Introduction Plant Breeding Systems

sexual asexual

panmixis(xenogamy)

self-fertilization

vegetative agamospermy

Page 11: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Reproductive strategies options

Introduction Richards 1997

Page 12: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Reproductive strategies options

Introduction de Jong & Klinkhamer 2005; Richards, 1997

72% 7% 4%7%

4% 5%

Page 13: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary
Page 14: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual theory

Sexual theory Plant Breeding Systems

'paradox of sex': Why so widespraed when it is unsure and costly?

Page 15: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual theory

Sexual theory Plant Breeding Systems

sexuality generates variability:

● recombination

● segregation

● syngamy

sexuality allows gene migration (spreading of successful mutation)

Page 16: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual theory

Sexual theory Antonovics & Ellstrand 1984, Evolution

'Tangled Bank' hypothesis (Felsenstein 1974) based on 'frequency dependent selection', reduced intragenotype competition

Sexuality is advantageous in heterogeneous environment

Page 17: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual theory

Sexual theory van Valen 1973, EvolTheor

'Red Queen' hypothesis (van Valen 1973) based on 'frequency dependent selection', biotic – biotic interactions

Sexuality is advantageous in heterogeneous environment

advantage of sex at the level of individuals, and the constant evolutionary arms race between competing species (co-evolution)

Page 18: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual theory

Sexual theory

1. cost of sex (cost of meiosis) = production of some portion of unfit progeny

Disadvantages of sexuality (compared to asexuality)

2. unsure reproduction (harsh environment and limited cross-pollination)

3. cost of sex (production of male gametes)

Page 19: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual reproduction in flowering plants

Sexual reproduction

1. no need of water for successful reproduction (vs ferns)

Evolutionary success of angiosperms (ca 300,000 species)

over ferns (ca 12,000) and gymnosperms (700 species)

2. protection and dispersion of zygote within seeds / fruits

3. wide range of pollination syndroms (vs gymnosperms)

4. wide range of growth forms (ecological succes) (vs gymnosperms)

Page 20: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual reproduction in flowering plants

Sexual reproduction

Female function: gynoecium (pistils)

Page 21: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual reproduction in flowering plants

Sexual reproduction

Male function: androecium (stamens)

Page 22: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual reproduction in flowering plants

Sexual reproduction

Double fecondation (fusion)

Page 23: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Sexual reproduction in flowering plants

Sexual reproduction Endress, 2011 AmJBot

Evolutionary trends in ovaries and ovules

Page 24: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Floral diversity and pollination

Sexual reproduction

Floral evolution

Page 25: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Floral diversity and pollination

Scent attraction and nectar reward

Pollination biology Knudsen 1999, Oikos

Page 26: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Floral diversity and pollination

Sexual reproduction

Pollination (zoo-, anemo-, hydro-)

Page 27: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Pollination biology and gene flow

Pollinator behaviour and foraging strategies

Pollination biology Schmitt 1980, Evolution; Kromer, 2008, PlantBiol

Page 28: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Floral diversity and pollination

Pollination biology Ghazoul 2006, JEcol

Page 29: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

hermaphrodites 95% (Richards 1997)

potential for selfing

40% can do, 20% usually

(ii) Why outcrossing is more frequent than self-fertilization?

Page 30: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

How do plants avoid self-polination?

Mechanisms:

mechanistic

temporal

genetic

Page 31: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mechanistic separation of male and female functions

allowing flower structures for cross-pollination:

cleistogamy vs chasmogamy

Page 32: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mechanistic separation of male and female functions

enlarging the distance between male and female structures

Page 33: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mechanistic separation of male and female functions

enlarging the distance between male and female structures

Dole 1992 AmJBot

Page 34: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Mechanistic separation of male and female functions -herkogamy (heterostyly)

Silvertown & Charlesworth 2007

Page 35: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Temporal separation of male and female functions -Dichogamy

A. protandry (male first)

Centaurea stoebe s.l.

Page 36: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Temporal separation of male and female functions -Dichogamy

A. protandry (male first)

MF

Centaurea stoebe s.l.

Page 37: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Temporal separation of male and female functions -Dichogamy

B. progyny (female first)

Magnolia grandiflora

Page 38: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Temporal separation of male and female functions -Dichogamy

Routley et al. 2004 IntJPlaSci

protandry

protogyny

Page 39: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Temporal separation of male and female functions -Dichogamy

Routley et al. 2004 IntJPlaSci

SI plants

SC plants

Page 40: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Temporal separation of male and female functions -Dichogamy

Routley et al. 2004 IntJPlaSci

SI plants

SC plants

Page 41: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

A. protandry - reducing the impact of pollen-pistil interference on pollen import and export (usually in SI taxa)

Temporal separation of male and female functions -Dichogamy

B. protogyny - reducing the negative impact of inbreeding (usually in SC taxa)

Page 42: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Genetic mechanism: multiallelic self-incompatibility

Self-incompatibility (SI) - the inability of fertile hermaphrodite plant to produces viable zygotes after self-pollination.

Genetically based (S-alleles, frequency dependent system)

Two types: Sporophytic SI and Gametophytic SI

Page 43: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Multiallelic self-incompatibility

Self-incompatibility

Gametophytic self-incompatibility (most of plants)

Page 44: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Multiallelic self-incompatibility

Sporophytic self-incompatibility (Asteraceae, Brassicaceae)

Self-incompatibility Hiscock & McInnis, 2008, TrendsPlantSciSelf-incompatibility

Page 45: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Multiallelic self-incompatibility

Break-down of SI system

Self-incompatibilitySelf-incompatibility

Page 46: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Floral heteromorphyphenomenon when 2 or 3 different floral morphs occur within population

mating compatible between morphs, incompatible within

Self-incompatibilitySelf-incompatibility

Page 47: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Dioecy

4% of flowering plants

more often in trees & shrubs than in perennial herbs

sometimes associated with 'sexual chromosomes'

Dioecy

Page 48: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Dioecy

Dioecy Charlesworth 2002, Heredity

Page 49: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Self-fertilization & inbreeding

Selfing

As ca 95% species are hermaphrodites > potential for selfing

Advantages?

Disadvantages?

Page 50: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Self-fertilization & inbreeding

Selfing Barrett 1998, TrendsPlantSci, Barrett et al 1996 PhilTrans

Page 51: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Agamospermy

Agamospermy Koltunow 2012, eLS

asexual seed production:progeny is a genetic copy of maternal plant

21 angiosperm families, Asteraceae, Poaceae & Rosaceae

associated with polyploidy, interspecific hybridization, SI (in sexual relatives), perenniality

21 angiosperm families, Asteraceae, Poaceae & Rosaceae

diplospory(Hieracium, Taracacum)

apospory(Pilosella, Rubus, Potentilla)

Page 52: Plant Breeding Systems - Univerzita Karlova · Sexual reproduction in flowering plants Sexual reproduction 1. no need of water for successful reproduction (vs ferns) Evolutionary

Agamospermy

Agamospermy

Advantages

Disadvantages