1 American Journal of Botany 89(11): OCCURRENCE AND PHYLOGENETIC SIGNIFICANCE OF LATEX IN THE MALPIGHIACEAE 1 ANDREA S. VEGA, 2,5 MARIA A. CASTRO, 3,5 AND WILLIAM R. ANDERSON 4,5 2 Instituto de Botánica Darwinion, Labardén 200, C. C. 22, B1642HYD Buenos Aires, Argentina; 3 Laboratorio de Anatomía Vegetal, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, C1428EHA Buenos Aires, Argentina; and 4 University of Michigan Herbarium, Suite 112, 3600 Varsity Drive, Ann Arbor, Michigan USA Latex and laticifers are reported for the first time in the genera Galphimia and Verrucularia (Malpighiaceae), with description and illustration of the leaf and stem anatomy of both genera. Those genera and the other two in which latex is known (Lophanthera and Spachea) constitute a single tribe, Galphimieae, that is at or near the base of the family s phylogeny, which suggests that latex in the Malpighiaceae may indicate an ancestor shared with the Euphorbiaceae. Key words: Galphimia brasiliensis; Galphimieae; latex; laticifers; Malpighiaceae; Verrucularia glaucophylla. The angiosperm family Malpighiaceae includes approximately 65 genera and 1250 species in the world and is distributed in tropical and subtropical regions of both hemispheres (W. R. Anderson, unpublished data). Nearly 950 species are endemic to the New World, northern South America being the major center of diversity (Anderson, 1979). The monophyly of the Malpighiaceae is supported by morphological characters (Anderson, 1979, 1990) and has recently been confirmed with molecular data (Chase et al., 1993; Cameron, Chase, and Anderson, 1995; Wurdack and Chase, 1996). Recent molecular studies of the Malpighiaceae did not provide convincing bases for dividing it into a small number of monophyletic subfamilies or tribes that reflect obvious morphological synapomorphies (Cameron et al., 2001; Davis, Anderson, and Donoghue, 2001), but they did confirm the monophyly of the tribe Galphimieae Nied. This tribe, as circumscribed by Anderson (1978), comprises four genera: Galphimia Cav., Lophanthera A. Juss., Spachea A. Juss., and Verrucularia A. Juss. Anderson (1981, 2001) mentioned the presence of milky latex in the stems of Lophanthera and Spachea, and the present paper documents the occurrence of latex in the other two genera, Galphimia and Verrucularia. Galphimia includes approximately ten species of tropical and subtropical America, with a disjunct distribution (Niedenzu, 1928). Most of the species grow in Mexico and Central America and only one, Galphimia brasiliensis (L.) A. Juss., is found in South America, in northeastern Argentina, southern Brazil, Bolivia, Paraguay, and Uruguay. This species has leaf and calyx glands, and the structural and ultrastructural anatomy of these glands was studied by Castro, Vega, and Múlgura (2001). The genus Verrucularia A. Juss. includes two species, V. glaucophylla A. Juss. and V. piresii W. R. Anderson. Verrucularia glaucophylla has only calyx glands and is endemic to Bahia, Brazil (Niedenzu, 1928). 1 Manuscript received 22 January 2002; revision accepted 21 May The authors thank André M. Amorim for his valuable help in collecting materials of Verrucularia glaucophylla and graphic designer Tomás E. Aversa for his technical assistance. 5 Authors for reprint requests ( fcen.uba.ar; MATERIALS AND METHODS Plant materials Fresh and fixed materials of Galphimia brasiliensis (L.) A. Juss. and Verrucularia glaucophylla A. Juss. were studied. Herbarium specimens were deposited in the herbarium of the Instituto de Botánica Darwinion (SI). Studied materials: Galphimia brasiliensis: ARGENTINA. Prov. Entre Ríos. Dpto. Colón: Parque Nacional El Palmar, Los Loros, 14/III/1998, Múlgura & Vega 1991; 4/XII/1998, Múlgura & Vega Dpto. Concordia: Parque Rivadavia, near José Hernández monument, 15/III/1998, Múlgura & Vega Verrucularia glaucophylla: BRAZIL. Bahia. Municipio de Morro do Chapéu, Cachoeira do Ferro Doido, ca m, 4/V/2001, Amorim et al (CEPEC, SI). Anatomical studies Leaves and stems of G. brasiliensis and V. glaucophylla were fixed in FAA (formalin : ethanol : acetic acid) and then stored in 70% ethanol. Part of each sample had been previously treated with sodium hypochlorite to eliminate contents. All materials were dehydrated in an ethanol series and embedded in paraffin after infiltration in a vacuum oven. Transverse and longitudinal sections 10 m thick were cut on a rotary microtome. Sections were double stained with safranin-fast green (D Ambrogio de Argüeso, 1986). Light microscope studies were made using a ZEISS Phomi III microscope (Zeiss, Oberkochen, Germany) and black and white Kodak Tmax 25 ASA film. RESULTS Field work During field work in Entre Ríos province (Argentina), the presence of latex in G. brasiliensis was observed. The latex is milky and was detected in stems and blades (Figs. 1 2). During the preparation of the present paper, Ch. Anderson (personal communication) told us about one collection of G. brasiliensis from Bolivia (Ortíz S. 7, MICH) whose label reported it to produce latex. Collectors of herbarium specimens of V. glaucophylla have never reported observing latex in the field, but laticifers were detected by anatomical techniques in stems and blades of that species. Anatomical studies Galphimia brasiliensis The blade in transverse and longitudinal sections is shown in Figs The adaxial epidermis is one-layered, the cells quadrangular to rectangular, with a thick outer tangential cell wall and cuticle. The papillate abaxial epidermis has a thin outer tangential cell wall. Trichomes are absent. Stomata are present in both surfaces and are more abundant in the abaxial epidermis. The 1725
2 1726 AMERICAN JOURNAL OF BOTANY [Vol. 89 Figs Galphimia brasiliensis (L.) A. Juss Field photographs of latex in petiole and blade, respectively. Figs Blade in transverse section, with laticifers immersed in spongy parenchyma. 3. Cleared section. Bar 50 m. 4. Uncleared section. Bar 50 m. 5. Blade in longitudinal section showing an articulated and unbranched laticifer. Bar 50 m. Figs Stem. 6. Transverse section. Bar 50 m. 7. Longitudinal section. Bar 50 m. ABE, abaxial epidermis; ADE, adaxial epidermis; B, blade; C, cuticle; CH, chlorenchyma; CO, cortex; L, laticifer; LX, latex; P, petiole; PH, phloem; PI, pith; PP, palisade parenchyma; S, stomata; SP, spongy parenchyma; VB, vascular bundles; X, xylem. dorsiventral mesophyll includes one layer of palisade parenchyma, composed of axially elongated cells slightly attached to one another and representing one-third of the total thickness. The spongy parenchyma has conspicuous intercellular spaces in which are immersed abundant laticifers distributed in the proximity of collateral vascular bundles. The laticifers were easily detected in transverse and longitudinal sections of the blades (Figs. 3 5) and stained intensively in uncleared sections (Fig. 4). In transverse section, laticifers are circular, ( 35.1) m in diameter, numerous, with a parallel and regular pattern in relation to the axis of the leaf and (51.3 ) m apart (Fig. 3 4). In longitudinal section, laticifers are tubular, articulated, not ramified, and not anastomosing (Fig. 5). The cells of the laticifers are m long, elongated, with dilated extremes like a trumpet and semi-thick terminal walls. The stem in transverse and longitudinal sections is illustrated in Figs The stem is circular in shape and is bounded by a one-layered epidermis with stomata. The cortex consists of subepidermal spongy chlorenchyma and parenchyma tissue, then secondary phloem and xylem and a conspicuous pith of parenchyma cells. Laticifers are distributed in the cortex and are scarcely recognizable due to their similarity to parenchyma cells (Fig. 7). Verrucularia glaucophylla The blade in transverse and longitudinal sections is shown in Figs The adaxial epidermis is one-layered, the cells quadrangular, with a thick outer tangential cell wall and cuticle. The conspicuously papillate abaxial epidermis exhibits one simple or occasionally bifurcate papilla on each cell, and a thick outer tangential cell wall. Trichomes are absent. Stomata are present in the abaxial epidermis overarched by adjacent papillate epidermal cells. The mesophyll is dorsiventral, including one-layered palisade parenchyma, which represents about one-half of the total thickness. The cells of typical palisade parenchyma are axially elongated and attached to one another. The spongy parenchyma has conspicuous intercellular spaces. Abundant laticifers are immersed between the palisade and spongy parenchyma, in the prox-
3 November 2002] VEGA ET AL. LATEX IN MALPIGHIACEAE 1727 Figs Verrucularia glaucophylla A. Juss. 8. Blade in transverse section showing laticifers immersed in the mesophyll. Bar 50 m. Figs Blade in paradermal section. 9. Bar 50 m. 10. Bar 100 m. Figs Stem in transverse section. 11. Bar 100 m. 12. Cuticle, epidermis, incipient periderm, and cortical parenchyma in detail. Bar 20 m. 13. Secondary xylem and central pith of parenchyma showing a peripheral starchy cell zone in detail. Bar 20 m. 14. Stem in longitudinal section showing cortical laticifers (arrow). Bar 25 m. ABE, abaxial epidermis; ADE, adaxial epidermis; C, cuticle; CO, cortex; F, phloem fibers; L, laticifers; PH, phloem; PE, periderm; PP, palisade parenchyma; S, stomata; SE, stem epidermis; SP, spongy parenchyma; ST, starchy parenchyma cells; SX, secondary xylem; T, trichomes.
4 1728 AMERICAN JOURNAL OF BOTANY [Vol. 89 imity of collateral vascular bundles. The laticifers were easily detected in both transverse and longitudinal sections of the blades (Figs. 8 10). In transverse section, the laticifers are circular, (31.2 ) ( 62.4) m in diameter, numerous, and (52 ) ( 176.8) m apart, with a parallel and irregular pattern in relation to the axis of the leaf (Fig. 8). In longitudinal section, the laticifers are tubular, articulated, not ramified, and not anastomosing. The cells of the laticifers are (369.2 ) ( 728) m long, uniform in diameter throughout their length (Fig. 9). In paradermal section the laticifers show a complex sinuous pattern (Fig. 10). The stem in transverse and longitudinal sections is illustrated in Figs The stem in early stages of secondary growth is circular in shape and surrounded by a one-layered epidermis with stomata (Fig. 11). The epidermal cells are radially elongated, with a thick outer tangential wall and cuticle. There is an incipient subepidermal periderm composed of 3 4 layers of cells (Fig. 12). The cortex contains 5 8 layers of parenchyma cells with laticifers and scarce starch and calcium oxalate druses. The vascular cylinder is composed of partially lignified phloem fibers, secondary phloem, and xylem. A central pith of parenchyma shows a peripheral zone of starchy cells (Fig. 13). Laticifers, present in the cortex, are easily recognized in longitudinal section (Fig. 14) but scarcely recognizable in transverse section because of their similarity to the parenchyma cells in which they are immersed. DISCUSSION Anderson (1978) restricted the tribe Galphimieae Nied. to four genera (Galphimia, Lophanthera, Spachea, and Verrucularia) on the basis of morphological characters. Cameron et al. (2001), using molecular data from the same four genera, found 100% bootstrap support for the tribe Galphimieae as restricted by Anderson, and Davis, Anderson, and Donoghue (2001) reported similar support, although their study lacked material of Verrucularia. Latex has previously been reported from only two genera of the Malpighiaceae, Lophanthera and Spachea (Anderson, 1981, 2001). The present study constitutes the first report of latex in Galphimia and Verrucularia and the first description to be published of the laticifers in this family. The fact that laticifers and latex are present in all four genera of the tribe Galphimieae provides a synapomorphy giving additional support to the monophyly of the tribe. There is an alternate, more interesting way to interpret the presence of latex in the tribe Galphimieae. The family Malpighiaceae was included in a large clade recently named Malpighiales (APG, 1998), with Violaceae, Passifloraceae, Linaceae, Clusiaceae, Peridiscaceae, and Euphorbiaceae, among others (Chase et al., 1993; Savolainen et al., 2000; Soltis et al., 2000). Anderson (1990) suggested that Lophanthera may be basal in the family, and the trees published recently (Cameron et al., 2001; Davis, Anderson, and Donoghue, 2001) place the tribe Galphimieae at or near the base of the family s phylogeny. It is therefore tempting to speculate that the presence of latex in this one tribe of Malpighiaceae constitutes a character linking the family to the Euphorbiaceae. In that interpretation, latex in the basal genera of the Malpighiaceae may be a symplesiomorphy inherited from a common ancestor shared with the Euphorbiaceae sensu stricto, i.e., the subfamilies Euphorbioideae and Crotonoideae of Webster (1975). Those are the groups of Euphorbiaceae that have laticifers, and they also share with the Malpighiaceae the character of having only one ovule in each locule of the ovary. The Euphorbioideae further resemble the Malpighiaceae in having simple leaves and simple hairs, their biglandular bracts recall the gland-bearing leaves and sepals of the Malpighiaceae, and their pollen is tricolporate, as in the basal clades of Malpighiaceae (Webster, 1975; Davis, Anderson, and Donoghue, 2001). The laticifers in the Euphorbioideae are non-articulated, whereas in the Crotonoideae, which in most characters are less like the Malpighiaceae, the laticifers are articulated like those reported here for the Galphimieae. Rudall (1987) argued that articulated and non-articulated laticifers probably evolved from a common ancestor in the Euphorbiaceae, and the evidence from the Malpighiaceae would seem to support the presence of articulated laticifers in the common ancestor of Euphorbiaceae sensu stricto and Malpighiaceae. Because the resolution of the phylogeny at the base of the Malpighiaceae is still rather labile, it would be best to regard the above phylogenetic speculations as preliminary, but they do point the way to future studies. It would be worthwhile to seek laticifers in the genera identified in the recent molecular studies (Cameron et al., 2001; Davis, Anderson, and Donoghue, 2001) as most closely related to the Galphimieae: Acmanthera Griseb., Blepharandra Griseb., Byrsonima L. C. Rich. ex Kunth, Coleostachys A. Juss., Diacidia Griseb., and Pterandra A. Juss. The fact that latex is unknown in those genera does not mean they do not have laticifers; latex had not been reported from Verrucularia, either, and we discovered its laticifers only because one of us (W. R. Anderson) predicted their existence on the basis of that genus s position in the tribe Galphimieae. If laticifers are basal in the Malpighiaceae, a symplesiomorphy shared with an outgroup like Euphorbiaceae, one would predict that laticifers, even if non-functional, may have been retained in some of the other genera near the base of the family s phylogeny. LITERATURE CITED ANDERSON, W. R [ 1977 ]. Byrsonimoideae, a new subfamily of the Malpighiaceae. Leandra 7: ANDERSON, W. R Floral conservatism in neotropical Malpighiaceae. Biotropica 11: ANDERSON, W. R Malpighiaceae. In The botany of the Guayana Highland: Part XI. Memoirs of the New York Botanical Garden 32: ANDERSON, W. R The origin of the Malpighiaceae the evidence from morphology. Memoirs of the New York Botanical Garden 64: ANDERSON, W. R Malpighiaceae. In P. E. Berry, K. Yatskievych, and B. K. Holst [eds.], Flora of the Venezuelan Guayana, vol. 6, Missouri Botanical Garden Press, St. Louis, Missouri, USA. APG [ANGIOSPERM PHYLOGENY GROUP] An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85: CAMERON, K. M., M. W. CHASE, AND W. R. ANDERSON Comparison of molecular trees from rbcl and matk analyses of Malpighiaceae. American Journal of Botany 82(supplement): 117 (Abstract). CAMERON, K. M., M. W. CHASE, W.R.ANDERSON, AND H. G. HILLS Molecular systematics of Malpighiaceae: evidence from plastid rbcl and matk sequences. American Journal of Botany 88: CASTRO, M. A., A. S. VEGA, AND M. E. MÚLGURA Structure and ultrastructure of leaf and calyx glands in Galphimia brasiliensis (Malpighiaceae). American Journal of Botany 88: CHASE, M. W., ET AL Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcl. Annals of the Missouri Botanical Garden 80: D AMBROGIO DE ARGÜESO, A Manual de Técnicas en Histología Vegetal. Hemisferio Sur S.A., Buenos Aires, Argentina.
5 November 2002] VEGA ET AL. LATEX IN MALPIGHIACEAE 1729 DAVIS, C. C., W. R. ANDERSON, AND M. J. DONOGHUE Phylogeny of Malpighiaceae: evidence from chloroplast ndhf and trnl-f nucleotide sequences. American Journal of Botany 88: NIEDENZU, F Malpighiaceae. In A. Engler [ed.], Das Pflanzenreich IV, 141. W. Engelmann, Leipzig, Germany. RUDALL, P. J Laticifers in Euphorbiaceae a conspectus. Botanical Journal of the Linnean Society 94: SAVOLAINEN, V., M. W. CHASE, S.B.HOOT, C.M.MORTON, D.E.SOLTIS, C. BAYER, M. F. FAY, A. Y. DE BRUIJN, S. SULLIVAN, AND Y. L. QIU Phylogenetics of flowering plants based upon a combined analysis of plastid atpb and rbcl gene sequences. Systematic Biology 49: SOLTIS, D.E.,ET AL Angiosperm phylogeny inferred from 18S rdna, rbcl, and atpb sequences. Botanical Journal of the Linnean Society 133: WEBSTER, G. L Conspectus of a new classification of the Euphorbiaceae. Taxon 24: WURDACK, K. J., AND M. W. CHASE Molecular systematics of Euphorbiaceae sensu lato using rbcl sequence data. American Journal of Botany 83(supplement): 203 (Abstract).
Department of Pharmacognosy 1, School of Pharmacy, Shaheed Beheshti University of Medical Sciences, and Faculty of Pharmacy 2, Tehran University of Medical Sciences, Tehran, Iran Anatomical study on Vaccinium
Lab Exercise 7: Leaves (also see Atlas pp. 141-150) In most green plants, leaves are the primary photosynthetic organs. They are well adapted for efficient light absorption, carbon fixation, and conduction
Plant Structure, Growth, & Development Ch. 35 Plants have organs composed of different tissues, which in turn are composed of different cell types A tissue is a group of cells consisting of one or more
Lecture 7: Plant Structure and Function I. Background A. Challenges for terrestrial plants 1. Habitat is divided a. Air is the source of CO2 for photosynthesis i. Sunlight cannot penetrate soil b. Soil
Plant Structure and Growth Plant body divided into root and shoot Shoot consists of leaves, buds, flowers, and stem Root consists of primary and secondary (lateral) roots Growth occurs at meristems Apical
Lecture 9 Outline (Ch. 35) I. Overview of plant systems II. III. IV. Major plant cell types Three plant tissues Three plant organs V. Plant growth A. Meristems B. Primary vs secondary C. Cell elongation
CELERY LAB - Structure and Function of a Plant READ ALL INSTRUCTIONS BEFORE BEGINNING! YOU MAY WORK WITH A PARTNER ON THIS ACTIVITY, BUT YOU MUST COMPLETE YOUR OWN LAB SHEET! Look at the back of this paper
Period Date LAB. LEAF STRUCTURE Plants are incredible organisms! They can make all their own food from the simple inputs of: sunlight air (carbon dioxide) water minerals This biological wizardry is accomplished
Plant Anatomy Objectives Identify and describe structure and function of each cell type and tissue type Describe organization of tissues and cells in each plant organ Relate function of an organ to structure
Plant Structure and Function Structure and Function Q: How are cells, tissues, and organs organized into systems that carry out the basic functions of a seed plant? 23.1 How are plant tissues organized?
Name Class Date Chapter 22 Plant Diversity Section Review 22-1 Reviewing Key Concepts Short Answer On the lines provided, answer the following questions. 1. Describe the main characteristics of plants.
CELERY LAB - Structure and Function of a Plant READ ALL INSTRUCTIONS BEFORE BEGINNING! YOU MAY WORK WITH A PARTNER ON THIS ACTIVITY, BUT YOU MUST COMPLETE YOUR OWN LAB SHEET! Plants are incredible organisms!
STRUCTURAL ORGANIZATION OF VASCULAR PLANTS STRUCTURAL ORGANIZATION OF VASCULAR PLANTS The vascular plant body plan Root system Shoot system Stems Leaves Flowers 2 PLANT TISSUES Tissue = collection of similar
Lab: Plant tissue systems and cell types In this lab we will become familiar with the main types of plant cells and tissues. You ll look at cells in the ground tissue, dermal tissue and vascular tissue.
LAB 10: PLANT ADAPTATIONS In this lab, you will 1. Identify the origin of modified structures and the environmental pressures that have led to the evolution of these adaptations (in the greenhouse with
Seed Plants: Gymnosperms and Angiosperms Answer the questions as you go through the power point, there are also paragraphs to read where you will need to hi-lite or underline as you read. 1. What are the
I. Leaf Structure and Anatomy Anatomy and Physiology of Leaves A. Structural Features of the Leaf Question: How do plants respire? Plants must take in CO 2 from the atmosphere in order to photosynthesize.
The Flower - what is it?! Floral structure will be examined in lab this Mon/Tues save space in your notes! Magnoliophyta - Flowering Plants! Introduction to Angiosperms "angio-" = vessel; so "angiosperm"
Biology SAT II Review Sheet Plants Plant Classification, Structure, Growth and Hormones Multicellular autotrophs (organisms that use the energy of inorganic materials to produce organic materials) Utilize
Araştırma Makalesi/Research Article A Morphological Study On Endemic Malabaila lasiocarpa Boiss. (Apiaceae) From Bingol (Turkey) Ömer KILIÇ 1 Abstract- In this study morphological characters of Malabaila
Lec 2. Plant body: form & function 1. Seed plants are the most successful land plants. Why? Main stages of a plant s life cycle. 2. Plants have developed appropriate structures to carry out the functions:
10 LESSON Leaf Structure and Transpiration INTRODUCTION Have you wondered what happens to all that water that disappears from the reservoir of your growing system? Although some might have evaporated from
Photosynthesis Light Energy transduction Chemical Energy (e.g. glucose) - Only photosynthetic organisms can do this (e.g. plants) - They are the ultimate source of chemical energy for all living organisms:
PHARMACOGNOSTICAL AND ANATOMICAL STUDIES OF LITSEA GLUTINOSA (LOUR.) C.B. ROB AN IMPORTANT MEDICINAL PLANT IN CHITTOOR DISTRICT OF ANDHRA PRADESH K. Tulasi Rao, M. Mahendra Nath*, K. Madhava Chetty Department
TRANSPORT IN PLANTS A. Introduction 1. The leaf is the main photosynthetic factory (Fig. 36.1, p. 702) a. This requires a transport system to move water and minerals from the roots to the leaf. This is
RESEARCH ARTICLE www.ijapc.com e-issn 2350-0204 Comparative Micromorphological Microscopic including Micrometric Evaluation of Important Tephrosia species - Leaves Urvi Ashani 1 * and Harisha C R 2 1,2
Exchange & Transport in Vascular Plants Plant Exchange & Transport I. II. Local Transport III. Long Distance Transport IV. Gas Exchange Water and Solutes - Uptake by Cells Passive Transport (Diffusion)
Digitization of the Albion College Herbarium Matthew Kleinow What is a Herbarium? Reference collection of dried or liquidpreserved plant specimens World s largest herbarium is at National Museum of Natural
10B Plant Systems Guided Practice Reproduction Station 1 1. Observe Plant A. Locate the following parts of the flower: stamen, stigma, style, ovary. 2. Draw and label the parts of a flower (listed above)
The Toledo Zoo/ThinkingWorks Teacher Overview for the Conservatory Lessons Teacher Overview: Conservatory Plants have many traits that are unique to this particular kingdom of living things. Below is a
Exercise 7 Angiosperm Reproduction: Flowers and Fruits Biol 1012, S2008, Lee, Etterson, and Little Goals Relate structures in a flower to the plant life cycle: alternation of generations. Identify floral,
No. 3 461 RINGS OF COLLAPSED CELLS IN PINUS RADIATA STEMWOOD FROM LYSIMETER-GROWN TREES SUBJECTED TO DROUGHT J. R. BARNETT Forest Research Institute, New Zealand Forest Service, Rotorua (Received for publication
Plant structure and function Have you ever considered where your food comes from? Fruits and vegetables come from plants, but what about meat, fats, dairy products and junk food? Meat comes from animals
TOPIC 5: SEEDLESS VASCULAR PLANTS (CH. 29) I. Vascular Plants (overview) plants with xylem and phloem 7 or 9 living phyla, depending on who you talk to able to dominate most terrestrial habitats because
Marine Angiosperms There are a wide variety of marine angiosperms some, such an the mangals, saltmarsh grasses and seagrasses that will either partly or entirely submerged (or submerged dependent on the
NCERT Solutions for TISSUES Question 1: What is a tissue? Tissue is a group of cells that are similar in structure and are organised together to perform a specific task. Question 2: What is the utility
from long shoots from a male tree and from a juvenile tree of still unknown sex exhibited normal stomata. The brachyblast leaves of the same trees, however, are Acta Botanica Neerlandica 12 (1963) 281-286
Laboratory 4 Plant Structure 2 Laboratory 4: Plant Structure OBJECTIVES After completing this lab you will be able to: 1. Differentiate between dicots and monocots within the following categories: a. root
3 How Plants Are Identified T he easiest method, and the one probably used the most often for learning the names of the plants of a particular location or region, or just the name of a particular plant,
Biology 172L General Biology Lab II Lab 03: Plant Life Cycles and Adaptations II: Gymnosperms and Angiosperms Introduction Vascular seed-bearing plants, such as gymnosperms (cone-bearing plants) and angiosperms
Lab 2- Bio 201 Prokaryotic and Eukaryotic Cells Name: OBJECTIVES To explore cell structure and morphology in prokaryotes and eukaryotes. To gain more experience using the microscope, and in particular,
1 of 30 Xylem tissue forms a continuous set of tubes that runs from the roots through stems and out into the spongy mesophyll of leaves. Active transport and root pressure cause water to move from soil
ISOLATION AND PROPERTIES OF SECRETORY GRANULES FROM RAT ISLETS OF LANGERHANS II Ultrastructure of the Beta Granule MARIE H GREIDER, S L HOWELL, and P E LACY From the Department of Pathology, Washington
Unit 1 Transport in plants A knowledge of how plants take in and transport substances is vitally important to agriculture. Using this knowledge, scientists are able to develop more effective ways of applying
BOTANY BASICS or Aren t plants absolutely amazing? By Donna Rea, Klamath County Master Gardener Plant characteristics All Plants Many celled not microscopic Cell walls surround cells Reproduce with spores
Celery Challenge: Investigating Water Movement in Plants Student s Guide In this module, you will think about transpiration, osmosis, and different cell types by trying to cause celery stalks to bend as
CHEM 107 Hair handout. 3-22-05 and 3-24-05 Basic Structure of Hair A hair can be defined as a slender, thread-like outgrowth from a follicle in the skin of mammals. Composed mainly of keratin, it has three
Systematics of hardy Theaceae and genus Stewartia Koen Camelbeke Arboretum Wespelaar Angiosperm Phylogeny Website Angiosperm Phylogeny Website Ericales Systematics Theaceae Family name Theaceae s.s. (versus
Biology 3B Laboratory Land Plant Structure Objectives Learn and recognize the basic tissues and structures of land plants Understand how these morphological features relate to plant structure and function
LAB 21 Using Bioinformatics to Investigate Evolutionary Relationships; Have a BLAST! Introduction: Between 1990-2003, scientists working on an international research project known as the Human Genome Project,
Lab 2- Bio 160 Name: Prokaryotic and Eukaryotic Cells OBJECTIVES To explore cell structure and morphology in prokaryotes and eukaryotes. To gain more experience using the microscope. To obtain a better
FABIAN ARMANDO MICHELANGELI Institute of Systematic Botany The New York Botanical Garden 200 th St. and Southern Blvd. Bronx, NY 10458 (last updated April 2007) e-mail: firstname.lastname@example.org Phone: (718) 817-8199
ACTIVITY 3 Introduction Several species of plant which grow in the Squares show clear morphological differences between leaves which grow in full sunlight and leaves which grow in the shade. In this investigation,
CURRICULUM VITAE ALEJANDRA VASCO Institute of Systematic Botany The New York Botanical Garden 200th St. & Kazimiroff Boulevard Bronx, New York 10458 Telephone: (1) 718.817.8102 E-mail: email@example.com
Lab Exercise Transport in Plants Objectives - Become familiar and be able to recognize the different types of cells found in the plant s vascular tissue. - Be able to describe root pressure and transpiration
Guidelines for Rare Plant Surveys Edited by Diana Bizecki Robson INTRODUCTION With the recent protection of some of Saskatchewan s rare plants under The Wildlife Act, industry will be required to conduct
Cells Cells are the fundamental unit of life. All living things are composed of cells. While there are several characteristics that are common to all cells, such as the presence of a cell membrane, cytoplasm,
BIO 311 Plant Structure and Development Lab Manual Dr. Alison Roberts Department of Biological Sciences University of Rhode Island Fall 2015 CONTENTS Laboratory schedule... 2 Lab 1 Introduction to plant
Teacher packs in Experimental Science Bio Pack 5 Examining flower structure Pack contents: A. Teachers Guide B. Students Guide C. Assessment Student s sheet D. Extensions to experiment E. Links to other
American Journal of Botany 91(1): 10 23. 2004. TREE FERN GROWTH STRATEGY IN THE LATE DEVONIAN CLADOXYLOPSID SPECIES PIETZSCHIA LEVIS FROM THE STUDY OF ITS STEM AND ROOT SYSTEM 1 AUDE SORIA 2 AND BRIGITTE
Tree Physiology and Growth Dr. Gary Chastagner Washington State University Puyallup, WA 98371 firstname.lastname@example.org March 2008 Principal Parts of a Vascular Plant http://extension.oregonstate.edu/mg/botany/external.html
WHAT ARE THE DIFFERENCES BETWEEN VASCULAR AND NON- VASCULAR PLANTS? Let s take a closer look. What makes them different on the outside and inside? Learning Intentions To understand how vascular plant cells
SEEDS A seed is a dormant embryonic plant. All seeds have several features in common: SEED COAT This is the outer layer of the seed. Function: Prevent destruction of the seed by dehydration or predation
Plant Diversity Unit 10- Plants /Study Guide KEY Answer Key SECTION 20.1. ORIGINS OF PLANT LIFE 1. eukaryotic, photosynthetic, same types of chlorophyll, starch as storage product, cellulose in cell walls
The outstanding and most significant feature of the flowering plants (and that which sets them out from other vascular plants) is the flower. Understanding the flower structure and names of the parts is
Bot. Wu and Bull. Kuo-Huang Acad. Sin. (1997) Calcium 38: crystals 97-104 in Moraceae 97 Calcium crystals in the leaves of some species of Moraceae Chi-Chih Wu and Ling-Long Kuo-Huang 1 Department of Botany,
3 levels of transport occur in plants: 1. Uptake of water and solutes by individual cells -for photosynthesis and respiration -ex: absorption of H 2 O /minerals by root hairs 2. Short distance cell-to-cell
THE CYTOCHEMICAL LOCALIZATION OF ASCORBIC ACID IN ROOT TIP CELLS BY WILLIAM A. JENSEN,* PH.D., AND LEROY G. KAVALJ-IAN,$ PH.D. (From the Kerckhoff Laboratories of Biology, California Institute of Technology,
The Huntington Library, Art Collections, and Botanical Gardens The Plant Cell: Peeping into Potatoes, Peppers, and Pears Adapted from Vodopich and Moore. Botany Laboratory Manual, WCB McGraw-Hill, 1998.
1 Topic 5. The Plant Cell Introduction: Cells are the fundamental units of life. Structurally, nothing simpler than a cell is actually alive. Plant cells, like animal cells (and unlike bacteria), are eukaryotic.
DNA Barcoding in Plants: Biodiversity Identification and Discovery University of Sao Paulo December 2009 W. John Kress Department of Botany National Museum of Natural History Smithsonian Institution New
Wood Sample Preparation for Microscopic Analysis Peter Prislan, Jožica Gričar, Katarina Čufar March 2014 COST is supported by the EU Framework Programme Horizon 2020 COST Action FP1106 Protocol for preparation
APPEARANCE GRADES NOVEMBER 2012 This document contains information on the primary grades of appearance timber. Appearance grades of timber, rough or surfaced two faces, shall unless otherwise specified,
A Correlation of Pearson To the Utah Core State Standards Resource Title: Publisher: Pearson Education publishing as Prentice Hall ISBN (10 or 13 digit unique identifier is required): SE: 9780133242003
Protein Sequence Analysis - Overview - UDEL Workshop Raja Mazumder Research Associate Professor, Department of Biochemistry and Molecular Biology Georgetown University Medical Center Topics Why do protein
Chapter 24 Reproduction of Seed Plants Section 24 1 Reproduction With Cones and Flowers (pages 609 616) Key Concepts What are the reproductive structures of gymnosperms and angiosperms? How does pollination
Operation Flower Dissection Classroom Activity: K-4 Time: One to two 50-minute class periods Overview: In this activity, students will observe the similarities and differences between flowers of different