Origin and Early Evolution of Chloroplasts
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Plant Evolution an Introduction to the History of Life
Plant Evolution An Introduction to the History of Life KARL J. NIKLAS The University of Chicago Press Chicago and London CONTENTS Preface vii Introduction 1 1 Origins and Early Events 29 2 The Invasion of Land and Air 93 3 Population Genetics, Adaptation, and Evolution 153 4 Development and Evolution 217 5 Speciation and Microevolution 271 6 Macroevolution 325 7 The Evolution of Multicellularity 377 8 Biophysics and Evolution 431 9 Ecology and Evolution 483 Glossary 537 Index 547 v Introduction The unpredictable and the predetermined unfold together to make everything the way it is. It’s how nature creates itself, on every scale, the snowflake and the snowstorm. — TOM STOPPARD, Arcadia, Act 1, Scene 4 (1993) Much has been written about evolution from the perspective of the history and biology of animals, but significantly less has been writ- ten about the evolutionary biology of plants. Zoocentricism in the biological literature is understandable to some extent because we are after all animals and not plants and because our self- interest is not entirely egotistical, since no biologist can deny the fact that animals have played significant and important roles as the actors on the stage of evolution come and go. The nearly romantic fascination with di- nosaurs and what caused their extinction is understandable, even though we should be equally fascinated with the monarchs of the Carboniferous, the tree lycopods and calamites, and with what caused their extinction (fig. 0.1). Yet, it must be understood that plants are as fascinating as animals, and that they are just as important to the study of biology in general and to understanding evolutionary theory in particular. -
Construct a Plant Evolution Timeline
Constructing a Timeline of Plant Evolution Use this photo as a guide to creating an interactive timeline. Refer to Geologic Time Scale charts. A good resource is http://www.ucmp.berkeley.edu/help/timeform.php Use the “Plant Timeline Cards” found at the end of this document. Attach pictures with Velcro to allow for manipulation by students, and to “build” the timeline in increments. Teachers will need to find and print photos for “seed ferns” or omit that step in the timeline. Cut and paste the text below under each picture. Cyanobacteria: Also known as blue- Mosses: These have the first identifiable green algae, these are bacteria transport systems for water (xylem), and (prokaryotes – no nucleus) that can reproduce via spores, which encase sex photosynthesize. cells to prevent desiccation. They still need to live near water. First eukaryotes (cell nucleus): Scientist think that eukaryotes evolved via Hornworts: These developed the cuticle symbiosis, with one bacteria engulfing (waxy coating to prevent water loss) and another, but not digesting it. stomata (openings in the cuticle to allow Chloroplasts were originally for gas exchange.) All land plants except cyanobacteria engulfed by another liverworts have these two features. bacterium. Development of a true vascular system Anabaena: These water organisms allowed for better transport of water evolved from being unicellular to more and provided structural support for the complex, multicellular organisms. Their plant to grow taller. specialized cells help with nitrogen fixation. Club mosses: These were the first plants to have true leaves with veins. They also Stoneworts: Scientists think that developed roots as an anchoring system colonization of land occurred only once, to allow plant to grow taller. -
The Big Bloom—How Flowering Plants Changed the World
The Big Bloom—How Flowering Plants Changed the World Written by Michael Klesius Republished from the pages of National Geographic magazine -- July 2002 In the summer of 1973 sunflowers appeared in my father's vegetable garden. They seemed to sprout overnight in a few rows he had lent that year to new neighbors from California. Only six years old at the time, I was at first put off by these garish plants. Such strange and vibrant flowers seemed out of place among the respectable beans, peppers, spinach, and other vegetables we had always grown. Gradually, however, the brilliance of the sunflowers won me over. Their fiery halos relieved the green monotone that by late summer ruled the garden. I marveled at birds that clung upside down to the shaggy, gold disks, wings fluttering, looting the seeds. Sunflowers defined flowers for me that summer and changed my view of the world. Flowers have a way of doing that. They began changing the way the world looked almost as soon as they appeared on Earth about 130 million years ago, during the Cretaceous period. That's relatively recent in geologic time: If all Earth's history were compressed into an hour, flowering plants would exist for only the last 90 seconds. But once they took firm root about 100 million years ago, they swiftly diversified in an explosion of varieties that established most of the flowering plant families of the modern world. Today flowering plant species outnumber by twenty to one those of ferns and cone-bearing trees, or conifers, which had thrived for 200 million years before the first bloom appeared. -
The Origin of Alternation of Generations in Land Plants
Theoriginof alternation of generations inlandplants: afocuson matrotrophy andhexose transport Linda K.E.Graham and LeeW .Wilcox Department of Botany,University of Wisconsin, 430Lincoln Drive, Madison,WI 53706, USA (lkgraham@facsta¡.wisc .edu ) Alifehistory involving alternation of two developmentally associated, multicellular generations (sporophyteand gametophyte) is anautapomorphy of embryophytes (bryophytes + vascularplants) . Microfossil dataindicate that Mid ^Late Ordovicianland plants possessed such alifecycle, and that the originof alternationof generationspreceded this date.Molecular phylogenetic data unambiguously relate charophyceangreen algae to the ancestryof monophyletic embryophytes, and identify bryophytes as early-divergentland plants. Comparison of reproduction in charophyceans and bryophytes suggests that the followingstages occurredduring evolutionary origin of embryophytic alternation of generations: (i) originof oogamy;(ii) retention ofeggsand zygotes on the parentalthallus; (iii) originof matrotrophy (regulatedtransfer ofnutritional and morphogenetic solutes fromparental cells tothe nextgeneration); (iv)origin of a multicellularsporophyte generation ;and(v) origin of non-£ agellate, walled spores. Oogamy,egg/zygoteretention andmatrotrophy characterize at least some moderncharophyceans, and arepostulated to represent pre-adaptativefeatures inherited byembryophytes from ancestral charophyceans.Matrotrophy is hypothesizedto have preceded originof the multicellularsporophytes of plants,and to represent acritical innovation.Molecular -
Cell Wall Ribosomes Nucleus Chloroplast Cytoplasm
Cell Wall Ribosomes Nucleus Nickname: Protector Nickname: Protein Maker Nickname: Brain The cell wall is the outer covering of a Plant cell. It is Ribosomes read the recipe from the The nucleus is the largest organelle in a cell. The a strong and stiff and made of DNA and use this recipe to make nucleus directs all activity in the cell. It also controls cellulose. It supports and protects the plant cell by proteins. The nucleus tells the the growth and reproduction of the cell. holding it upright. It ribosomes which proteins to make. In humans, the nucleus contains 46 chromosomes allows water, oxygen and carbon dioxide to pass in out They are found in both plant and which are the instructions for all the activities in your of plant cell. animal cells. In a cell they can be found cell and body. floating around in the cytoplasm or attached to the endoplasmic reticulum. Chloroplast Cytoplasm Endoplasmic Reticulum Nickname: Oven Nickname: Gel Nickname: Highway Chloroplasts are oval structures that that contain a green Cytoplasm is the gel like fluid inside a The endoplasmic reticulum (ER) is the transportation pigment called chlorophyll. This allows plants to make cell. The organelles are floating around in center for the cell. The ER is like the conveyor belt, you their own food through the process of photosynthesis. this fluid. would see at a supermarket, except instead of moving your groceries it moves proteins from one part of the cell Chloroplasts are necessary for photosynthesis, the food to another. The Endoplasmic Reticulum looks like a making process, to occur. -
Brown Algae and 4) the Oomycetes (Water Molds)
Protista Classification Excavata The kingdom Protista (in the five kingdom system) contains mostly unicellular eukaryotes. This taxonomic grouping is polyphyletic and based only Alveolates on cellular structure and life styles not on any molecular evidence. Using molecular biology and detailed comparison of cell structure, scientists are now beginning to see evolutionary SAR Stramenopila history in the protists. The ongoing changes in the protest phylogeny are rapidly changing with each new piece of evidence. The following classification suggests 4 “supergroups” within the Rhizaria original Protista kingdom and the taxonomy is still being worked out. This lab is looking at one current hypothesis shown on the right. Some of the organisms are grouped together because Archaeplastida of very strong support and others are controversial. It is important to focus on the characteristics of each clade which explains why they are grouped together. This lab will only look at the groups that Amoebozoans were once included in the Protista kingdom and the other groups (higher plants, fungi, and animals) will be Unikonta examined in future labs. Opisthokonts Protista Classification Excavata Starting with the four “Supergroups”, we will divide the rest into different levels called clades. A Clade is defined as a group of Alveolates biological taxa (as species) that includes all descendants of one common ancestor. Too simplify this process, we have included a cladogram we will be using throughout the SAR Stramenopila course. We will divide or expand parts of the cladogram to emphasize evolutionary relationships. For the protists, we will divide Rhizaria the supergroups into smaller clades assigning them artificial numbers (clade1, clade2, clade3) to establish a grouping at a specific level. -
(SSC) Region of Chloroplast Genomes1
NEWS & VIEWS AMERICAN JOURNAL OF BOTANY LETTER TO THE EDITOR Sources of inversion variation in the small single copy (SSC) region of chloroplast genomes1 Joseph F. Walker 2 , Robert K. Jansen 3,4 , Michael J. Zanis 5 , and Nancy C. Emery6,7 Modern sequencing technology has led to a proliferation of whole- Walker et al., 2014 ; Zhang et al., 2014 ; Wang et al., 2015 ). Th ese genome sequences of chloroplasts in a growing number of plant analyses compare the SSC orientation among lineages using a single lineages, bringing opportunities for comparisons that provide in- plastome to represent each lineage and thus have missed the within- sights into the evolutionary history of the plastomes and their host individual variation that exists in this region. Currently, whole- plants ( Jansen et al., 2007 ; Doorduin et al., 2011 ). Amid the emerg- chloroplast genomes are published in GenBank without preference ing literature in this area is a hypothesis that the small single copy for the orientation of the SSC region, leading to apparent variation (SSC) region is a “hotspot” for inversion events (sensu Liu et al., in the orientation of the SSC region among individuals that is actu- 2013 ) because diff erent orientations of the region have been re- ally due to chloroplast heteroplasmy within individuals ( Wolfe and ported in relatively high frequencies among closely related taxa Randle, 2004 ), as originally described by Palmer (1983) . For exam- ( Liu et al., 2013 ; Walker et al., 2014 ). We would like to draw atten- ple, two sequences of Lactuca sativa that have been independently tion to a study by Palmer (1983) that bears heavily on this discus- published (NC_007578 and DQ_383816) were entered with diff er- sion, yet has been overlooked by several authors of publications ent orientations of the SSC region, which could be interpreted as investigating whole-chloroplast genome sequence order, including a major inversion existing within the species if the investigators are one study by some of the authors of this letter ( Walker et al., 2014 ). -
Evolution of Land Plants P
Chapter 4. The evolutionary classification of land plants The evolutionary classification of land plants Land plants evolved from a group of green algae, possibly as early as 500–600 million years ago. Their closest living relatives in the algal realm are a group of freshwater algae known as stoneworts or Charophyta. According to the fossil record, the charophytes' growth form has changed little since the divergence of lineages, so we know that early land plants evolved from a branched, filamentous alga dwelling in shallow fresh water, perhaps at the edge of seasonally-desiccating pools. The biggest challenge that early land plants had to face ca. 500 million years ago was surviving in dry, non-submerged environments. Algae extract nutrients and light from the water that surrounds them. Those few algae that anchor themselves to the bottom of the waterbody do so to prevent being carried away by currents, but do not extract resources from the underlying substrate. Nutrients such as nitrogen and phosphorus, together with CO2 and sunlight, are all taken by the algae from the surrounding waters. Land plants, in contrast, must extract nutrients from the ground and capture CO2 and sunlight from the atmosphere. The first terrestrial plants were very similar to modern mosses and liverworts, in a group called Bryophytes (from Greek bryos=moss, and phyton=plants; hence “moss-like plants”). They possessed little root-like hairs called rhizoids, which collected nutrients from the ground. Like their algal ancestors, they could not withstand prolonged desiccation and restricted their life cycle to shaded, damp habitats, or, in some cases, evolved the ability to completely dry-out, putting their metabolism on hold and reviving when more water arrived, as in the modern “resurrection plants” (Selaginella). -
The Structure and Function of Plastids
The Structure and Function of Plastids Edited by Robert R. Wise University of Wisconsin, Oshkosh WI, USA and J. Kenneth Hoober Arizona State University, Tempe AZ, USA Contents From the Series Editor v Contents xi Preface xix A Dedication to Pioneers of Research on Chloroplast Structure xxi Color Plates xxxiii Section I Plastid Origin and Development 1 The Diversity of Plastid Form and Function 3–25 Robert R. Wise Summary 3 I. Introduction 4 II. The Plastid Family 5 III. Chloroplasts and their Specializations 13 IV. Concluding Remarks 20 Acknowledgements 21 References 21 2 Chloroplast Development: Whence and Whither 27–51 J. Kenneth Hoober Summary 27 I. Introduction 28 II. Brief Review of Plastid Evolution 28 III. Development of the Chloroplast 32 IV. Overview of Photosynthesis 43 References 46 3 Protein Import Into Chloroplasts: Who, When, and How? 53–74 Ute C. Vothknecht and J¨urgen Soll Summary 53 I. Introduction 54 II. On the Road to the Chloroplast 56 III. Protein Translocation via Toc and Tic 58 IV. Variations on Toc and Tic Translocation 63 V. Protein Translocation and Chloroplast Biogenesis 64 VI. The Evolutionary Origin of Toc and Tic 66 VII. Intraplastidal Transport 66 VIII. Protein Translocation into Complex Plastids 69 References 70 xi 4 Origin and Evolution of Plastids: Genomic View on the Unification and Diversity of Plastids 75–102 Naoki Sato Summary 76 I. Introduction: Unification and Diversity 76 II. Endosymbiotic Origin of Plastids: The Major Unifying Principle 78 III. Origin and Evolution of Plastid Diversity 85 IV. Conclusion: Opposing Principles in the Evolution of Plastids 97 Acknowledgements 98 References 98 5 The Mechanism of Plastid Division: The Structure and Origin of The Plastid Division Apparatus 103–121 Shin-ya Miyagishima and Tsuneyoshi Kuroiwa Summary 104 I. -
Gene Genealogies and Population Variation in Plants
Colloquium Gene genealogies and population variation in plants Barbara A. Schaal* and Kenneth M. Olsen Department of Biology, Washington University, St. Louis, MO 63130 Early in the development of plant evolutionary biology, genetic selection. Morphology, karyotypes, and fitness components are drift, fluctuations in population size, and isolation were identified central traits for understanding evolution and adaptation, but as critical processes that affect the course of evolution in plant they limit which evolutionary processes can be studied. In his species. Attempts to assess these processes in natural populations book, Stebbins discusses such events as fluctuations in popula- became possible only with the development of neutral genetic tion size, random fixation (genetic drift), and isolation as all markers in the 1960s. More recently, the application of historically affecting the process of evolution (see passage quoted above; ref. ordered neutral molecular variation (within the conceptual frame- 1). However, the study of these mechanisms requires markers work of coalescent theory) has allowed a reevaluation of these that are not under selection. microevolutionary processes. Gene genealogies trace the evolu- In the years after the publication of Stebbins’ book, among tionary relationships among haplotypes (alleles) with populations. the first major technical advances in evolutionary biology were Processes such as selection, fluctuation in population size, and the development of protein electrophoresis and the identifi- population substructuring affect the geographical and genealog- cation of allozyme variation in natural populations. Many ical relationships among these alleles. Therefore, examination of allozymes are selectively neutral, and thus, for the first time, these genealogical data can provide insights into the evolutionary evolutionary biologists could attempt to assess the amount of history of a species. -
Algal Chloroplasts Secondary Article
Algal Chloroplasts Secondary article Saul Purton, University College London, London, UK Article Contents . Introduction A great diversity of chloroplasts is found amongst the various algal groups. This diversity . Diversity and Classification of Algae is the result of an intriguing evolutionary process that involved the acquisition of . Origins and Evolution of Algal Chloroplasts chloroplasts by different eukaryotic organisms. Chloroplast Genetics and Molecular Biology . Protein Transport in the Chloroplast . Summary Introduction The chloroplast is one of a family of related biosynthetic and which lack the differentiated structures that organelles (termed plastids) found within the cells of define higher plants (roots, shoots, leaves, etc.). Indeed, plants, eukaryotic algae and certain protists. The primary the algae are often referred to as ‘lower’ or ‘primitive’ role of the chloroplast is the fixation of atmospheric carbon plants. Included within the algae are the prokaryotic through photosynthesis, but it is also the site of synthesis of cyanobacteria (formerly referred to as blue-green algae), many other important compounds including pigments, together with a diverse collection of microscopic and fatty acids, amino acids and nucleotides. Chloroplasts are macroscopic eukaryotes. Algal species can be unicellular, distinguishable from other plastid types in that they filamentous or multicellular and they range in size from the contain chlorophyll and other pigments that are involved unicellular forms that are only a few micrometres in in light energy capture and dissipation. In higher plants, diameter to the giant Laminaria seaweeds that are tens of nonphotosynthetic plastids such as chromoplasts, amylo- metres long. Algae have adapted to life in a wide range of plasts and leucoplasts are found in nongreen tissue and environments. -
Chloroplasts Are the Food Producers of the Cell. the Organelles Are Only Found in Plant Cells and Some Protists Such As Algae
Name: ___________________________ Cell #2 H.W. due September 22nd, 2016 Period: _________ Chloroplasts are the food producers of the cell. The organelles are only found in plant cells and some protists such as algae. Animal cells do not have chloroplasts. Chloroplasts work to convert light energy of the Sun into sugars that can be used by cells. It is like a solar panel that changes sunlight energy into electric energy. The entire process is called photosynthesis and it all depends on the little green chlorophyll molecules in each chloroplast. In the process of photosynthesis, plants create sugars and release oxygen (O2). The oxygen released by the chloroplasts is the same oxygen you breathe every day. Chloroplasts are found in plant cells, but not in animal cells. The purpose of the chloroplast is to make sugars that feed the cell’s machinery. Photosynthesis is the process of a plant taking energy from the Sun and creating sugars. When the energy from the Sun hits a chloroplast and the chlorophyll molecules, light energy is converted into the chemical energy. Plants use water, carbon dioxide, and sunlight to make sugar and oxygen. During photosynthesis radiant energy or solar energy or light energy is transferred into chemical energy in the form of sugar (glucose). You already know that during photosynthesis plants make their own food. The food that the plant makes is in the form of sugar that is used to provide energy for the plant. The extra sugar that the plant does not use is stored as starch for later use. Mitochondria are known as the powerhouses of the cell.