Evolution of Plants David Baum Game plan • What are “plants” and how did they evolve? • Differences between plant and animal evolution • Some stories of plant evolution What are the three most important* events in the evolution of life on earth? 1. Oxygenic photosynthesis (cyanobacteria) 2. Invasion of land (plants) 3. Human agriculture and technology *Profoundly affecting the globe’s chemistry and ecology Early land plants were low to the ground They became larger, more complex, and acquired a vascular system Time Multiple origins of “trees” Crane and Leslie (2013) Why? http://www.earthhistory.org.uk/recolonisation/vegetation-in-devonian An evolutionary arm’s race • The Red Queen principle Now, here, I see it takes all the running you can do, to keep in the same place. If you want to get somewhere else, you must run at least twice as fast as that! (Through the Looking Glass, Lewis Carroll) Competition for light is a very important driver of plant evolution Problems that plants faced • Gain light, water, nutrients • Escaping predators (once animals invaded land) • Sex! If you want to know more: Botany 130, 300, 305, 401, 500 Fern sperm cell Are there differences between plant and animal evolution? • Very few – plants are excellent “model systems” • But.. – Greater diversity in sexual systems • Abundant asexuality – More chemistry less behavior – Maybe more evolution by “hopeful monsters” Examples of “hopeful monsters?” Rudall PJ, Bateman RM. 2003. Trends Plant Sci. 8(2):76-82. Rudall PJ, Bateman RM. 2002. Biol Rev Camb Philos Soc.77(3):403-441 Are flowers monsters? Pollen cone Living gymnosperms have unisexual cones Seed cone Are flowers monsters? Megasporophyll Microsporophyll (stamen) A flower is a bisexual “cone” (although unisexual flowers have evolved in many groups) If so: quite successful! • ~300,000 species of flowering plants • Dominate all land ecosystems (and several aquatic ones) • Provide all food resources for humans Are there differences between plant and animal evolution? • Very few – plants are excellent “model systems” • But.. – Greater diversity in sexual systems • Abundant asexuality – More chemistry less behavior – Maybe more evolution by “hopeful monsters” – Polyploidy Why is polyploidy common in plants? • Tolerance of different ploidy levels during development (dosage compensation?) • Often make unreduced gametes (“rescues” meiotic problems) Chester et al. 2010. Genes 1(2), 166-192. Why is polyploidy common in plants? • Tolerance of different ploidy levels during development (dosage compensation?) • Often make unreduced gametes (“rescues” meiotic problems) • Allows for “instant speciation” http://www.amjbot.org/content/95/6/713/F1.large.jpg Example of “instant speciation” • Spartina maritima grew in British salt marshes • A different species, S. alterniflora arrived ~1816, probably from boats arriving from America – This was figured out partly by talking to thatchers in the 1830’s who noted that the new form of cordgrass had appeared some 20 years earlier Example of “instant speciation” • In 1879 two botanist brothers, H. & J. Groves, found a plant that confused them: – “Spartina which . . .would be rather more S. alterniflora… yet which we now consider to be [S. maritima]” • In 1880 they concluded they had found a new species, which they called S. townsendii • It was not fertile • Examination of the herbarium records showed that it had appeared by 1870 Example of “instant speciation” • Fertile “S. townsendii” appeared and was eventually named as a new species S. anglica. • Stapf (1910) noted: ‘towards the end of the eighties (c. 1890) something occurred that favoured the spreading of the grass’ – this is now presumed to represent the origin of S. anglica. • Stapf (1926) hypothesized that S. anglica and S. townsendii are hybrids of S. maritima X S. alterniflora Example of “instant speciation” • S. anglica spread quickly around Southampton harbor • It gradually reached other estuaries – especially during World War II – perhaps because of training with landing craft http://www.thinkdefence.co.u k/2011/11/uk-militarybridging-world-war-ii-africaand-northwest-europe/ Example of “instant speciation” • But if this is true: how come “…it is fertile and breeds true to type” (Hutchins, 1930)? [S. anglica] Hypothesis S. maritima 2a chromosomes S. townsendii a+ b chromosomes (sterile) S. alterniflora 2b chromosomes Hypothesis S. townsendii a+ b chromosomes (sterile) Chromosome doubling S. anglica 2a + 2b This hypothesis explains a few things • Why the S. townsendii arose suddenly • Why it was successful (“hybrid vigor”) • Why it was sterile • Why S. anglica arose suddenly • Why it was successful • Why it is fertile But is it true? C. Leonard Huskins (1930) Professor of Genetics at McGill S. maritima, 2n = 56 (“2a”) S. alterniflora, 2n = 70 (“2b”) 126 S. anglica, 2n = (Became Professor of Botany at UWMadison in 1945) Huskins (1930) Genetica There are many other examples of polyploid species in plants • Often generate competitively superior species • Many crops have arisen by hybrid polyploidy (including wheat and potato) • Means that plants have lots of “extra” genes that can acquire new function • All vascular plants have undergone multiple rounds of polyploidy in their ancestry Pollination Stories Pollination (only occurs in seed plants) avoids the need for motile sperm • Pollen is a minute male plant • Can be carried by wind (rarely water) • More commonly animals do it – Insects – Birds – Mammals Pollen needs to deliver the gametes to the egg cells Stigma Pollen tubes grow through plant tissues – navigated chemically Pollen tubes grow through plant tissues – navigated chemically Plants have evolved diverse ways to get pollen from one flower to another • Wind • Water (rare) • Animals – Mutualistic (give a reward) – Parasitic (trick the animal) How do you think this evolved? • What else would you like to know? At the other extreme: Figs and fig wasps figs are “tomb blossoms” Implications • There is a one-to-one relationship between a fig species and its wasp pollinator species • Predicts cospeciation: that the figs and wasp Prediction: • One-to-one species association • Cospeciation A a Fig phylogeny C c B b D d E e F f Wasp phylogeny Actual result Host switching Weiblen and Bush (2002) • Plant evolution is similar to other multicellular eukaryotes • Arm’s race for light • Polyploidy is especially important • Coevolution with animals for pollination (and dispersal) is important • (Also, abundant coevolution with fungi and bacteria) Feel free to contact me: dbaum@wisc.edu