The Role of Tissue Cell Polarity in Monocot Development

Total Page:16

File Type:pdf, Size:1020Kb

The Role of Tissue Cell Polarity in Monocot Development The Role of Tissue Cell Polarity in Monocot Development Annis Richardson Thesis submitted for the Degree of Doctor of Philosophy University of East Anglia John Innes Centre September 2015 © This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that use of any information derived there from must be in accordance with current UK Copyright Law. In addition, any quotation or extract must include full attribution. Abstract Nature exhibits huge diversity in organ shape, and yet all organs start as small bud-like peripheral outgrowths. Combinations of different spatial and temporal developmental switches in shape determine final organ shape. In plants shape arises through growth, which is defined by axiality and growth rates. Here I tested three hypotheses for how developmental switches in shape could arise: (1) growth rates alone are altered, (2) axiality alone is altered (3) both growth rates and axiality are altered. Using a multidisciplinary approach I explored which of the hypotheses was true for developmental switches in shape during organ development in two monocot models: early grass leaf development and the Hooded barley mutant. Developmental switches in shape were first volumetrically described using 3D imaging. Using this framework, computational models were generated to formulate hypotheses which could account for the switches in shape. Model predictions were then tested using whole-mount immunolocalisation of SISTER OF PINFORMED 1 (SoPIN1), gene expression, and cell division and shape analyses. Synthetic biology was also used to generate a set of transgenic tools for future testing of the models. I found that a developmental switch in shape during early grass leaf development may arise through alterations in growth rates alone (hypothesis 1). In contrast, ectopic flower and wing formation in Hooded may arise through modulation of growth rates and axiality combined (hypothesis 2). In this case a single gene, BKn3, triggers the growth change, possibly through directly influencing tissue cell polarity (if axiality is defined by a polarity based axiality system), with differential effects on shape depending on where it is expressed. This suggests that novel developmental switches in shape could evolve due to single gene mutations, and that during evolution, modulation of growth may have been redeployed in different spatial and temporal patterns to trigger novel changes in shape, ultimately changing final form. 2 Contents Abstract ......................................................................................................................... 2 Contents ........................................................................................................................ 3 List of Tables .................................................................................................................. 8 List of Figures ................................................................................................................. 9 Acknowledgements ...................................................................................................... 13 Introduction ................................................................................................................. 15 1.1 Developmental switches ...................................................................... 15 1.2 Growth in plant tissues ........................................................................ 16 1.3 Growth and developmental switches in shape ...................................... 18 1.4 Axiality systems ................................................................................... 19 1.4.1 Polarity based axiality system ......................................................... 19 1.4.2 Stress based axiality system ............................................................ 23 1.4.3 Markers of axiality: PINs and hairs ................................................. 24 1.5 The contribution of different tissues within the organ ........................... 25 1.6 Computational models at different scales ............................................. 26 1.7 Exploring dicot and monocot development ........................................... 27 1.8 This work ............................................................................................. 31 2 Grass Leaf Development ................................................................................... 32 2.1 Leaf development in the grasses ........................................................... 32 2.2 Aim of this project ................................................................................ 37 2.3 Describing a developmental switch in shape during primordial grass leaf development ................................................................................................... 37 2.4 The formation of the hood from a ring primordium could be accounted for by enhanced anisotropic growth towards the midvein ................................. 50 2.5 Axial information in the early grass leaf primordium ............................. 55 2.6 A change in axial information and/ or growth rate pattern may be required for the next shape transition from a hood to a cone ............................ 60 2.7 Axial information is oriented towards the midvein tip after the hood stage consistent with both model predictions ................................................... 66 2.8 Exploring growth rate patterns across the grass leaf primordium .......... 69 2.9 Discussion ............................................................................................ 77 2.9.1 Characterising developmental switches in shape during primordial stages of grass leaf development ................................................................ 77 2.9.2 Modelling the primordial stages of grass leaf development .......... 77 3 2.9.3 Predicted changes in growth during grass leaf development ......... 78 2.9.4 The role of growth rate patterns in grass leaf development .......... 79 2.9.5 The role of axial information in grass leaf development ................ 80 2.9.6 Insights into the evolution of the grass leaf .................................... 81 2.9.7 Future work and concluding remarks .............................................. 82 3 How Can a Single Gene Induce a Developmental Switch in Shape? The Hooded Barley Mutant .............................................................................................................. 85 3.1 Barley floral development and the Hooded mutant ............................... 85 3.1.1 Morphology of wild-type barley ...................................................... 85 3.1.2 Morphology of the Hooded barley mutant ..................................... 87 3.1.3 Previous studies in the Hooded mutant. ......................................... 90 3.2 Aim of this project ............................................................................... 92 3.3 Characterising a developmental switch in shape in the barley flower: Staging ectopic flower development ................................................................. 92 3.4 Ectopic expression of BKn3 in the Hooded lemma precedes the formation of the ectopic meristem ................................................................................. 104 3.5 The ectopic expression of BKn3 in the Hooded lemma induces a reorientation in axial information at the cellular level ..................................... 110 3.6 The ectopic expression of BKn3 in the Hooded lemma induces changes in the expression pattern of candidate polarity organisers .................................. 123 3.7 Specific changes in growth in the lemma margin trigger the developmental switch in shape responsible for wing formation ...................... 136 3.7.1 Characterising a possible second developmental switch in shape in the Hooded mutant .................................................................................... 137 3.7.2 Modelling Hooded lemma wing development as a consequence of changes in growth ...................................................................................... 140 3.7.3 Axial information may specifically reorient at the margin of the Hooded lemma ........................................................................................... 144 3.7.4 BKn3 may act cell autonomously in the margins to form the wings in the Hooded mutant ................................................................................ 148 3.8 Discussion .......................................................................................... 151 3.8.1 Characterising the morphology of the wild-type and Hooded spike during development ................................................................................... 151 3.8.2 There may be two independent developmental switches in shape in the Hooded mutant .................................................................................... 152 3.8.3 Single genes are able to trigger developmental switches in shape through modulating growth rates and axiality .......................................... 153 4 3.8.4 BKn3 may influence axial information through modulating the expression of organiser components ......................................................... 154 3.8.5 Future work and concluding remarks ............................................ 156 4 Developing a Transgenic Toolkit in Barley ......................................................
Recommended publications
  • Annual Report 2015
    Annual Report 2015 Xishuangbanna Tropical Botanical Garden Chinese Academy of Sciences Cover photos, anti-clockwise: 1. Prunus kunmingensis T. Su, P. Wilf et Z.K. Zhou, (a–e) KUN PC2015001-KUN PC2015005. (f) CT scan showing longitudinal section and seed (P. 11); 2. 2015 Annual Conference of Chinese Botanic Gardens (P. 25); 3. UK Prince William visited XTBG (P. 55); 4. The launching ceremony of the renovation program of CAS-SEABRI (P. 5); 5. The international "Fascination of Plants Day 2015" (P. 38); 6. XTBG researchers assessed the current status of all orchid species in Xishuangbanna (P. 9); 7. Ecologists and conservation biologists change thinking by Anthropocene concept (P. 9). Annual Report 2015 Xishuangbanna Tropical Botanical Garden Chinese Academy of Sciences March 29, 2016 Xishuangbanna Tropical Botanical Garden (XTBG), Chinese Academy of Sciences is a non-profit, comprehensive botanical garden involved in scientific research, plant diversity conservation XTBG’s vision: and public science education, affiliated directly to Desirable base for plant diversity conservation the Chinese Academy of Sciences. and ecological studies. Noah’s Ark for tropical plants. XTBG’s mission: Promote science development and environmental conservation through implementing scientific research on ecology and plant diversity conservation, horticultural exhibition, and public education. Photo by XIAO Jiao-Jiao CONTENTS SCIENCE .............................................................................. 2 Project Development ...............................................................
    [Show full text]
  • The Relationship Between Gene Network Structure and Expression Variation Among Individuals and Species
    UC San Diego UC San Diego Previously Published Works Title The Relationship between Gene Network Structure and Expression Variation among Individuals and Species. Permalink https://escholarship.org/uc/item/77v6f7dc Journal PLoS genetics, 11(8) ISSN 1553-7390 Authors Sears, Karen E Maier, Jennifer A Rivas-Astroza, Marcelo et al. Publication Date 2015-08-28 DOI 10.1371/journal.pgen.1005398 Peer reviewed eScholarship.org Powered by the California Digital Library University of California RESEARCH ARTICLE The Relationship between Gene Network Structure and Expression Variation among Individuals and Species Karen E. Sears1,2*, Jennifer A. Maier1, Marcelo Rivas-Astroza3, Rachel Poe4, Sheng Zhong3, Kari Kosog1, Jonathan D. Marcot1, Richard R. Behringer5, Chris J. Cretekos6†, John J. Rasweiler, IV7, Zoi Rapti4 1 School of Integrative Biology, University of Illinois, Urbana, Illinois, United States of America, 2 Institute for Genomic Biology, University of Illinois, Urbana, Illinois, United States of America, 3 Department of Bioengineering, University of California, San Diego, La Jolla, California, United States of America, 4 Department of Mathematics, University of Illinois, Urbana, Illinois, United States of America, 5 Department of Genetics, University of Texas MD Anderson Cancer Center, Houston, Texas, United States of America, 6 Department of Biological Sciences, Idaho State University, Pocatello, Idaho, United States of America, 7 Department of Obstetrics and Gynecology, State University of New York Downstate Medical Center, Brooklyn, New York, United States of America † Deceased. * [email protected] OPEN ACCESS Citation: Sears KE, Maier JA, Rivas-Astroza M, Poe R, Zhong S, Kosog K, et al. (2015) The Relationship between Gene Network Structure and Expression Abstract Variation among Individuals and Species.
    [Show full text]
  • Download Bamboo Records (Public Information)
    Status Date Accession Number Names::PlantName Names::CommonName Names::Synonym Names::Family No. Remaining Garden Area ###########2012.0256P Sirochloa parvifolia Poaceae 1 African Garden ###########1989.0217P Thamnocalamus tessellatus mountain BamBoo; "BergBamBoes" in South Africa Poaceae 1 African Garden ###########2000.0025P Aulonemia fulgor Poaceae BamBoo Garden ###########1983.0072P BamBusa Beecheyana Beechy BamBoo Sinocalamus Beechyana Poaceae 1 BamBoo Garden ###########2003.1070P BamBusa Burmanica Poaceae 1 BamBoo Garden ###########2013.0144P BamBusa chungii White BamBoo, Tropical Blue BamBoo Poaceae 1 BamBoo Garden ###########2007.0019P BamBusa chungii var. BarBelatta BarBie BamBoo Poaceae 1 BamBoo Garden ###########1981.0471P BamBusa dolichoclada 'Stripe' Poaceae 2 BamBoo Garden ###########2001.0163D BamBusa dolichoclada 'Stripe' Poaceae 1 BamBoo Garden ###########2012.0069P BamBusa dolichoclada 'Stripe' Poaceae 1 BamBoo Garden ###########1981.0079P BamBusa dolichomerithalla 'Green Stripe' Green Stripe Blowgun BamBoo Poaceae 1 BamBoo Garden ###########1981.0084P BamBusa dolichomerithalla 'Green Stripe' Green Stripe Blowgun BamBoo Poaceae 1 BamBoo Garden ###########2000.0297P BamBusa dolichomerithalla 'Silverstripe' Blowpipe BamBoo 'Silverstripe' Poaceae 1 BamBoo Garden ###########2013.0090P BamBusa emeiensis 'Flavidovirens' Poaceae 1 BamBoo Garden ###########2011.0124P BamBusa emeiensis 'Viridiflavus' Poaceae 1 BamBoo Garden ###########1997.0152P BamBusa eutuldoides Poaceae 1 BamBoo Garden ###########2003.0158P BamBusa eutuldoides
    [Show full text]
  • Plants Unlimited Rufa Clump Bamboo
    [email protected] 207.594.7754 P.O. Box 374 629 Commercial St. Rockport, Maine 04856 Rufa Clump Bamboo* Fargesia rufa Height: 10 feet Spread: 8 feet Sunlight: Hardiness Zone: 4b Other Names: Fargesia dracocephala Description: This bamboo makes a beautiful screen or accent in the garden, incredibly architectural when mature; a non-invasive species; attractive sheaths are rusty red; a must for Oriental gardens Rufa Clump Bamboo Photo courtesy of NetPS Plant Finder Ornamental Features Rufa Clump Bamboo's narrow leaves remain green in color throughout the year. Neither the flowers nor the fruit are ornamentally significant. Landscape Attributes Rufa Clump Bamboo is an herbaceous evergreen perennial with an indistinguished habit of growth. Its relatively fine texture sets it apart from other garden plants with less refined foliage. This is a relatively low maintenance plant, and can be pruned at anytime. It has no significant negative characteristics. Rufa Clump Bamboo foliage Rufa Clump Bamboo is recommended for the following Photo courtesy of NetPS Plant Finder landscape applications; - Accent - Vertical Accent - Mass Planting - Hedges/Screening - General Garden Use - Naturalizing And Woodland Gardens Visit plants-unlimited.com [email protected] 207.594.7754 P.O. Box 374 629 Commercial St. Rockport, Maine 04856 Planting & Growing Rufa Clump Bamboo will grow to be about 10 feet tall at maturity, with a spread of 8 feet. Its foliage tends to remain dense right to the ground, not requiring facer plants in front. It grows at a fast rate, and under ideal conditions can be expected to live for approximately 20 years.
    [Show full text]
  • Over 1,000 Varieties of Edible Plants!
    OVER 1,00 0 VA RIE TI ES O F E D IB L E P L A N T S ! VISIT OUR NURSERY! 6469 SE 134th Ave, Portland, OR 97236 ORDER TOLL FREE (503) 208-7520 1-877-353-4028 | fax: 1-800-418-9983 One Green World is a family owned nursery and garden center located in Portland, Oregon. We provide a huge EMAIL [email protected] selection of fruiting trees, shrubs, berries, vines, unique citrus, nut trees, vegetables and much more to people GET THE LATEST NEWS all over the United States. In addition to the plants we Sign up for our e-mail newsletter online! sell, the experts at OGW are available for questions and www.onegreenworld.com/newsletter advice on plants, plant care and gardening techniques. We will share gardening tips and let you know about plant sales, special events, unique tastings and classes We specialize in all things edible and are continuously throughout the year. adventuring to discover new and unique plant varieties. Our vision is to create a One Green World where PRICING everyone has access to homegrown delicious and Prices are subject to change at anytime. Our most nutritious fruits and veggies. accurate pricing for all plants can be found online at www.OneGreenWorld.com HAVE QUESTIONS? We're here to help! PLANT INDEX ACACIAS ............................104 GIANT GROUNDCHERRY .....101 NECTARINES ......................72 TREE COLLARD ...................96 AGAVE ..............................108 GINGER ..............................94 OAK TREES .........................90 ULLUCO .............................2 ALMOND ...........................86
    [Show full text]
  • Transcriptomic and Epigenomic Characterization of the Developing Bat Wing
    ARTICLES OPEN Transcriptomic and epigenomic characterization of the developing bat wing Walter L Eckalbar1,2,9, Stephen A Schlebusch3,9, Mandy K Mason3, Zoe Gill3, Ash V Parker3, Betty M Booker1,2, Sierra Nishizaki1,2, Christiane Muswamba-Nday3, Elizabeth Terhune4,5, Kimberly A Nevonen4, Nadja Makki1,2, Tara Friedrich2,6, Julia E VanderMeer1,2, Katherine S Pollard2,6,7, Lucia Carbone4,8, Jeff D Wall2,7, Nicola Illing3 & Nadav Ahituv1,2 Bats are the only mammals capable of powered flight, but little is known about the genetic determinants that shape their wings. Here we generated a genome for Miniopterus natalensis and performed RNA-seq and ChIP-seq (H3K27ac and H3K27me3) analyses on its developing forelimb and hindlimb autopods at sequential embryonic stages to decipher the molecular events that underlie bat wing development. Over 7,000 genes and several long noncoding RNAs, including Tbx5-as1 and Hottip, were differentially expressed between forelimb and hindlimb, and across different stages. ChIP-seq analysis identified thousands of regions that are differentially modified in forelimb and hindlimb. Comparative genomics found 2,796 bat-accelerated regions within H3K27ac peaks, several of which cluster near limb-associated genes. Pathway analyses highlighted multiple ribosomal proteins and known limb patterning signaling pathways as differentially regulated and implicated increased forelimb mesenchymal condensation in differential growth. In combination, our work outlines multiple genetic components that likely contribute to bat wing formation, providing insights into this morphological innovation. The order Chiroptera, commonly known as bats, is the only group of To characterize the genetic differences that underlie divergence in mammals to have evolved the capability of flight.
    [Show full text]
  • Hortax Presentation
    HORTAX CULTIVATED PLANT TAXONOMY GROUP Muddles and misapplications: taxonomic difficulties associated with bamboos in cultivation Chris Stapleton bamboo-identification.co.uk formerly: University of Aberdeen Forestry Department Royal Botanic Garden Edinburgh Royal Botanic Gardens Kew Presentation roughly as given in 2015, updated a little in 2021. Here I illustrate the problem – multiple names in the UK for the same cultivated plant. Genus, species & cultivar names all varying on displayed and marketed plants. All these are the same species. To decide which name is correct Steve Renvoize & I had to go as far as conserving a species name in the journal Taxon. The genus Pleioblastus is considered appropriate on the grounds of morphology and molecular data. In that genus it is now correctly called Pleioblastus viridistriatus, but you will still see these other names around. Popping in to Tesco’s, I saw a nice little bamboo plant on the stands outside. It had an authoritative label, saying Fountain Bamboo. It is not Fountain Bamboo, which is Fargesia nitida. On the back it says Fargesia rufa, which it also is not. It is actually Fargesia dracocephala ‘Rufa’, Dragon-head Bamboo. It also says it comes from the mountains of western China, while in fact it comes from Central China, but that is not so important as getting the name so wrong. How have we got into this position in Europe? This disinformation would not happen in the US, where we have verified and standardised bamboo names for most of the horticulturally available bamboos. Stapleton, C. M. A. & Renvoize, S. A. (2001). Proposal to conserve the name Bambusa viridistriata Siebold ex André (Poaceae, Bambusoideae).
    [Show full text]
  • Genetic and Molecular Control of Vertebrate Limb Development and Morphological Diversity
    Genetic and Molecular Control of Vertebrate Limb Development and Morphological Diversity Lee Niswander HHMI/ U Colorado Health Sciences Center [email protected] Limb Development Chick Pro: experimental embryology, rapid functional analysis Con: relative lack of genetics Mouse Pro: genetics, genetic manipulation Con: in utero development Bat Pro: evo-devo studies, experimental embryology Con: relative lack of genetics relatively difficult to obtain Diversity of Diversity of Vertebrate Limbs Vertebrate Limbs A s= stylopod s Pr D z= zeugopod z a a= autopod P Modified from: Ridley M. Evolution, 2nd ed. A-P= Anterior-Posterior Blackwell Science 1996 Pr-D= Proximal-Distal P. Martin Int. J. Dev. Biol 1990 The thickened ectoderm at the distal rim of the limb bud is termed the apical ectodermal ridge. Laying down of the cartilage condensations E11.5 Martin, 1990 E12.5 Khokha et al. 2003 s z a Specification Elaboration of Patterning s z a 3 QuickTime™ and a 4 TIFF (LZW) decompressor 2 3 4 are needed to see this picture. 2 5 5 a z ssor ? ture. s pg pg S Z A Plzf/Gli3 dKO Barna et al. Fgf4/Fgf8 dKO Hoxa11/d11 dKO Sun et al. Nature (2002) Davis et al. Nature (1995) 1 Maria Barna AC D H Sin3A N-CoR/ SMRT POZ Zn+ Zn+ Zn+ Zn+ Zn+ Zn+ Zn+ Zn+ Zn+ PLZF (also known as ZFP 145) PLZF translocated to the Retinoic Acid Receptor Alpha (RARα)gene is associated with acute promyelocytic leukemia (APL) PLZF is a sequence specific DNA binding transcriptional repressor belonging to the POK (POZ and Kruppel) family of proteins that can remodel chromatin to a heterochromatic state.
    [Show full text]
  • 19493 OID Ctlg
    The Journal of the American Bamboo Society Volume 28 BAmBoo Science & culTure The Journal of the American Bamboo Society is published by the American Bamboo Society Copyright 2015 ISSN 0197– 3789 Bamboo Science and Culture: The Journal of the American Bamboo Society is the continuation of The Journal of the American Bamboo Society President of the Society Board of Directors Susanne Lucas David Benfield Bill Hollenback Vice President Carole Meckes Bill Hollenback Luc Boeraeve Karlyn M. Bradshaw Treasurer W.P. Edwards, III Dawn Weaver Susanne Lucas David King Secretary Dain Sansome David King John Tkach Danielo Ramos Membership Harry Simmons Kristen Comstock Membership Information Membership in the American Bamboo Society and one ABS chapter is for the calendar year and includes a subscription to the bimonthly Newsletter and annual Journal. See http://www.bamboo.org for current rates. Cover photo: Cover Photo: Phyllostachys in Prafrance (courtesy Patrick Cassiers) Bamboo Science and Culture: The Journal of the American Bamboo Society 28(1): 1-9 © Copyright 2015 by the American Bamboo Society Biogenic volatile organic compound emissions from bamboo: Exploring patterns of diversity across species Andrea N. Melnychenko and Todd N. Rosenstiel Department of Biology, Portland State University 1719 SW 10th Ave, Portland, OR 97201 USA ABSTRACT Emissions of biogenic volatile organic compounds (BVOCs) from plant leaves play significant roles in biological and atmospheric processes. BVOCs emissions can involve a diverse number of compounds and are an important method of plant signaling. However, some emissions of volatile compounds can negatively impact air quality at a regional scale. In order to better understand the role of BVOCs in plant physiology and chemical ecology, and to better predict how these emissions may alter air quality, the underlying relationships between these diverse compounds must be addressed.
    [Show full text]
  • 1 503-647-2700 Front Page- Need Hi
    front page- need Hi res closeup www.bamboogarden.com 1 503-647-2700 Forward by Ted Meredith Bamboo Garden is very special. Founded in 1984, Bamboo Garden has a diverse collection of more than 300 bamboo species and forms on 20 pastoral acres near North Plains, Oregon. Here mature bamboo groves are cast in a beautiful natural setting of rolling hills, ponds, mountain stream, and wooded backdrop. Customers of the nursery are offered golf cart tours of the extensive grounds. Europe has a number of splendid bamboo gardens, and a few very famous ones that are connected with a bamboo nursery, where one can see many species of bamboo in mature natural groves and then have the opportunity to purchase the same bamboos for one’s own garden. America now has an equivalent in the Bamboo Garden. Owner Ned Jaquith, an ardent bamboo enthusiast, has introduced countless people to the world of bamboo and served as a mentor to countless more---myself included. Nothing seems to please Ned more than introducing another person to bamboo. His welcom- ing enthusiasm carries through to the Bamboo Gar- den’s knowledgeable staff, who are adept at discuss- ing bamboos with experts and novices alike. Nursery manager Noah Bell oversees the operation, includ- ing nursery, office, and sales. Maintenance foreman Reveriano Ramirez directs bamboo propagation and care. Bamboo Garden is an impressive operation with many fine people in key roles working to make it so. Like Bamboo Garden, the Bamboo Garden catalog is something special too. The bamboos are beautifully photographed and described (Noah and Ned did most of the photography themselves) with clear illustra- tions that show how to maintain bamboo (credit to Charissa Brock for illustrations and layout design) .
    [Show full text]
  • Biological Sciences
    A Comprehensive Book on Environmentalism Table of Contents Chapter 1 - Introduction to Environmentalism Chapter 2 - Environmental Movement Chapter 3 - Conservation Movement Chapter 4 - Green Politics Chapter 5 - Environmental Movement in the United States Chapter 6 - Environmental Movement in New Zealand & Australia Chapter 7 - Free-Market Environmentalism Chapter 8 - Evangelical Environmentalism Chapter 9 -WT Timeline of History of Environmentalism _____________________ WORLD TECHNOLOGIES _____________________ A Comprehensive Book on Enzymes Table of Contents Chapter 1 - Introduction to Enzyme Chapter 2 - Cofactors Chapter 3 - Enzyme Kinetics Chapter 4 - Enzyme Inhibitor Chapter 5 - Enzymes Assay and Substrate WT _____________________ WORLD TECHNOLOGIES _____________________ A Comprehensive Introduction to Bioenergy Table of Contents Chapter 1 - Bioenergy Chapter 2 - Biomass Chapter 3 - Bioconversion of Biomass to Mixed Alcohol Fuels Chapter 4 - Thermal Depolymerization Chapter 5 - Wood Fuel Chapter 6 - Biomass Heating System Chapter 7 - Vegetable Oil Fuel Chapter 8 - Methanol Fuel Chapter 9 - Cellulosic Ethanol Chapter 10 - Butanol Fuel Chapter 11 - Algae Fuel Chapter 12 - Waste-to-energy and Renewable Fuels Chapter 13 WT- Food vs. Fuel _____________________ WORLD TECHNOLOGIES _____________________ A Comprehensive Introduction to Botany Table of Contents Chapter 1 - Botany Chapter 2 - History of Botany Chapter 3 - Paleobotany Chapter 4 - Flora Chapter 5 - Adventitiousness and Ampelography Chapter 6 - Chimera (Plant) and Evergreen Chapter
    [Show full text]
  • University of Cape Town Faculty of Science Department of Molecular and Cell Biology
    The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgementTown of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the Universityof of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University University of Cape Town Faculty of Science Department of Molecular and Cell Biology Limbs gone batty: Town The role of the anterior-posterior patterning signal, Sonic Hedgehog, in the Cape development of the unique bat limb. of Oorit Hockman Supervisors: Prof. Nicola II ling (MCI3) and Prof. David Jacobs (Zoology) University Research report Submitted in fulfillment of the requirements for the degree of Master of Science (Molecular Bio~gy) December 2007 "Twinkle, twinkle, little batl How I wonder what you're at! Up above the world you fly, Like a teatray in the sky." Town Cape of -The Mad Halter lewis Carroll's Alice's Adventures in Wonderland. University Table of Contents List of Figures and Tables IV Abbreviations vi Acknowledgements vii Abstract viii Chapter 1: Hypotheses for the molecular mechanisms of bat limb development 1.1. Introduction 1 1.2. The unique bat limb 1 1.3. Ecological hypotheses for bat wing development 3 1.4. The morphological development of the bat limb 3 1.5. The molecular mechanisms of limb development Town 5 1.5.1. Sonic Hedgehog and the patterning of the anterior-posterior axis 6 1.5.2. Activation and maintenance of the Sonic Hedgehog signal 8 1.5.3.
    [Show full text]