Plant Cell BIOBASE Corporation
HOME HELP FEEDBACK SUBSCRIPTIONS ARCHIVE SEARCH TABLE OF CONTENTS
 QUICK SEARCH:   [advanced]


     


First published online June 13, 2003; 10.1105/tpc.011544

This Article
Right arrow Full Text
Right arrow Full Text (PDF)
Right arrow Supplemental Data
Right arrow All Versions of this Article:
15/7/1538    most recent
tpc.011544v1
Right arrow Alert me when this article is cited
Right arrow Alert me if a correction is posted
Services
Right arrow Email this article to a friend
Right arrow Similar articles in this journal
Right arrow Similar articles in ISI Web of Science
Right arrow Similar articles in PubMed
Right arrow Alert me to new issues of the journal
Right arrow Download to citation manager
Right arrow reprints & permissions
Citing Articles
Right arrow Citing Articles via HighWire
Right arrow Citing Articles via CrossRef
Right arrow Citing Articles via ISI Web of Science (120)
Right arrow Citing Articles via Google Scholar
Google Scholar
Right arrow Articles by Parenicová, L.
Right arrow Articles by Colombo, L.
Right arrow Search for Related Content
PubMed
Right arrow PubMed Citation
Right arrow Articles by Parenicová, L.
Right arrow Articles by Colombo, L.
Agricola
Right arrow Articles by Parenicová, L.
Right arrow Articles by Colombo, L.
The Plant Cell, Vol. 15, 1538-1551, July 2003, Copyright © 2003,
American Society of Plant Biologists

Molecular and Phylogenetic Analyses of the Complete MADS-Box Transcription Factor Family in Arabidopsis

New Openings to the MADS World

Lucie Paenicová1,a, Stefan de Folter1,b, Martin Kieffer1,c, David S. Hornerd, Cristina Favallia, Jacqueline Busscherb, Holly E. Cookc, Richard M. Ingramc, Martin M. Katere, Brendan Daviesc, Gerco C. Angenentb and Lucia Colombo2,a

a Dipartimento di Biologia, Universitá degli Studi di Milano, 20133 Milan, Italy
b Plant Development and Reproduction, Plant Research International B.V., 6700 AA Wageningen, The Netherlands
c Centre for Plant Sciences, University of Leeds, Leeds LS2 9JT, United Kingdom
d Dipartimento di Fisiologia e Biochimica Generali, Universitá degli Studi di Milano, 20133 Milan, Italy
e Dipartimento di Genetica e di Biologia dei Microrganismi, Università degli Studi di Milano, 20133 Milan, Italy

2 To whom correspondence should be addressed. E-mail lucia.colombo{at}unimi.it; fax 39-02-50314764

MADS-box transcription factors are key regulators of several plant development processes. Analysis of the complete Arabidopsis genome sequence revealed 107 genes encoding MADS-box proteins, of which 84% are of unknown function. Here, we provide a complete overview of this family, describing the gene structure, gene expression, genome localization, protein motif organization, and phylogenetic relationship of each member. We have divided this transcription factor family into five groups (named MIKC, M{alpha}, M{beta}, M{gamma}, and M{delta}) based on the phylogenetic relationships of the conserved MADS-box domain. This study provides a solid base for functional genomics studies into this important family of plant regulatory genes, including the poorly characterized group of M-type MADS-box proteins. MADS-box genes also constitute an excellent system with which to study the evolution of complex gene families in higher plants.




This article has been cited by other articles:


Home page
jashsHome page
H. Yamane, Y. Kashiwa, T. Ooka, R. Tao, and K. Yonemori
Suppression Subtractive Hybridization and Differential Screening Reveals Endodormancy-associated Expression of an SVP/AGL24-type MADS-box Gene in Lateral Vegetative Buds of Japanese Apricot
J. Amer. Soc. Hort. Sci., September 1, 2008; 133(5): 708 - 716.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
J. G. Steffen, I.-H. Kang, M. F. Portereiko, A. Lloyd, and G. N. Drews
AGL61 Interacts with AGL80 and Is Required for Central Cell Development in Arabidopsis
Plant Physiology, September 1, 2008; 148(1): 259 - 268.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
M. Bemer, M. Wolters-Arts, U. Grossniklaus, and G. C. Angenent
The MADS Domain Protein DIANA Acts Together with AGAMOUS-LIKE80 to Specify the Central Cell in Arabidopsis Ovules
PLANT CELL, August 1, 2008; 20(8): 2088 - 2101.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
O. N. Danilevskaya, X. Meng, D. A. Selinger, S. Deschamps, P. Hermon, G. Vansant, R. Gupta, E. V. Ananiev, and M. G. Muszynski
Involvement of the MADS-Box Gene ZMM4 in Floral Induction and Inflorescence Development in Maize
Plant Physiology, August 1, 2008; 147(4): 2054 - 2069.
[Abstract] [Full Text] [PDF]


Home page
ANN BOT (LOND)Home page
A. T. Ku, Y.-S. Huang, Y.-S. Wang, D. Ma, and K.-W. Yeh
IbMADS1 (Ipomoea batatas MADS-box 1 gene) is Involved in Tuberous Root Initiation in Sweet Potato (Ipomoea batatas)
Ann. Bot., July 1, 2008; 102(1): 57 - 67.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
M. J. Carmona, J. Chaib, J. M. Martinez-Zapater, and M. R. Thomas
A molecular genetic perspective of reproductive development in grapevine
J. Exp. Bot., July 1, 2008; 59(10): 2579 - 2596.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
G. Wang, U. Ellendorff, B. Kemp, J. W. Mansfield, A. Forsyth, K. Mitchell, K. Bastas, C.-M. Liu, A. Woods-Tor, C. Zipfel, et al.
A Genome-Wide Functional Investigation into the Roles of Receptor-Like Proteins in Arabidopsis
Plant Physiology, June 1, 2008; 147(2): 503 - 517.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
C. H. Leseberg, C. L. Eissler, X. Wang, M. A. Johns, M. R. Duvall, and L. Mao
Interaction study of MADS-domain proteins in tomato
J. Exp. Bot., May 17, 2008; (2008) ern094v1.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
P. J. Rushton, M. T. Bokowiec, S. Han, H. Zhang, J. F. Brannock, X. Chen, T. W. Laudeman, and M. P. Timko
Tobacco Transcription Factors: Novel Insights into Transcriptional Regulation in the Solanaceae
Plant Physiology, May 1, 2008; 147(1): 280 - 295.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
S. Lee, Y.-M. Woo, S.-I. Ryu, Y.-D. Shin, W. T. Kim, K. Y. Park, I.-J. Lee, and G. An
Further Characterization of a Rice AGL12 Group MADS-Box Gene, OsMADS26
Plant Physiology, May 1, 2008; 147(1): 156 - 168.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
D. Thakare, W. Tang, K. Hill, and S. E. Perry
The MADS-Domain Transcriptional Regulator AGAMOUS-LIKE15 Promotes Somatic Embryo Development in Arabidopsis and Soybean
Plant Physiology, April 1, 2008; 146(4): 1663 - 1672.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
I.-H. Kang, J. G. Steffen, M. F. Portereiko, A. Lloyd, and G. N. Drews
The AGL62 MADS Domain Protein Regulates Cellularization during Endosperm Development in Arabidopsis
PLANT CELL, March 1, 2008; 20(3): 635 - 647.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
E. Piwarzyk, Y. Yang, and T. Jack
Conserved C-Terminal Motifs of the Arabidopsis Proteins APETALA3 and PISTILLATA Are Dispensable for Floral Organ Identity Function
Plant Physiology, December 1, 2007; 145(4): 1495 - 1505.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
J. M. Barrero, R. Gonzalez-Bayon, J. C. del Pozo, M. R. Ponce, and J. L. Micol
INCURVATA2 Encodes the Catalytic Subunit of DNA Polymerase {alpha} and Interacts with Genes Involved in Chromatin-Mediated Cellular Memory in Arabidopsis thaliana
PLANT CELL, September 1, 2007; 19(9): 2822 - 2838.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
C. Kutter, H. Schob, M. Stadler, F. Meins Jr., and A. Si-Ammour
MicroRNA-Mediated Regulation of Stomatal Development in Arabidopsis
PLANT CELL, August 1, 2007; 19(8): 2417 - 2429.
[Abstract] [Full Text] [PDF]


Home page
GeneticsHome page
P. A. Reeves, Y. He, R. J. Schmitz, R. M. Amasino, L. W. Panella, and C. M. Richards
Evolutionary Conservation of the FLOWERING LOCUS C-Mediated Vernalization Response: Evidence From the Sugar Beet (Beta vulgaris)
Genetics, May 1, 2007; 176(1): 295 - 307.
[Abstract] [Full Text] [PDF]


Home page
Mol Biol EvolHome page
A. S. Veron, K. Kaufmann, and E. Bornberg-Bauer
Evidence of Interaction Network Evolution by Whole-Genome Duplications: A Case Study in MADS-Box Proteins
Mol. Biol. Evol., March 1, 2007; 24(3): 670 - 678.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
W. Verelst, H. Saedler, and T. Munster
MIKC* MADS-Protein Complexes Bind Motifs Enriched in the Proximal Region of Late Pollen-Specific Arabidopsis Promoters
Plant Physiology, January 1, 2007; 143(1): 447 - 460.
[Abstract] [Full Text] [PDF]


Home page
Mol Biol EvolHome page
L. C. Hileman, J. F. Sundstrom, A. Litt, M. Chen, T. Shumba, and V. F. Irish
Molecular and Phylogenetic Analyses of the MADS-Box Gene Family in Tomato
Mol. Biol. Evol., November 1, 2006; 23(11): 2245 - 2258.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
T. H. Teeri, A. Uimari, M. Kotilainen, R. Laitinen, H. Help, P. Elomaa, and V. A. Albert
Reproductive meristem fates in Gerbera
J. Exp. Bot., October 1, 2006; 57(13): 3445 - 3455.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
R. Wang, K. Chong, and T. Wang
Divergence in spatial expression patterns and in response to stimuli of tandem-repeat paralogues encoding a novel class of proline-rich proteins in Oryza sativa
J. Exp. Bot., August 1, 2006; 57(11): 2887 - 2897.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
M. F. Portereiko, A. Lloyd, J. G. Steffen, J. A. Punwani, D. Otsuga, and G. N. Drews
AGL80 Is Required for Central Cell and Endosperm Development in Arabidopsis
PLANT CELL, August 1, 2006; 18(8): 1862 - 1872.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
C. P. Scutt, M. Vinauger-Douard, C. Fourquin, C. Finet, and C. Dumas
An evolutionary perspective on the regulation of carpel development
J. Exp. Bot., July 1, 2006; 57(10): 2143 - 2152.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
V. Gregis, A. Sessa, L. Colombo, and M. M. Kater
AGL24, SHORT VEGETATIVE PHASE, and APETALA1 Redundantly Control AGAMOUS during Early Stages of Flower Development in Arabidopsis
PLANT CELL, June 1, 2006; 18(6): 1373 - 1382.
[Abstract] [Full Text] [PDF]


Home page
Mol Biol EvolHome page
S. De Bodt, G. Theissen, and Y. Van de Peer
Promoter Analysis of MADS-Box Genes in Eudicots Through Phylogenetic Footprinting
Mol. Biol. Evol., June 1, 2006; 23(6): 1293 - 1303.
[Abstract] [Full Text] [PDF]


Home page
ANN BOT (LOND)Home page
P. WALCH-LIU, I. I. IVANOV, S. FILLEUR, Y. GAN, T. REMANS, and B. G. FORDE
Nitrogen Regulation of Root Branching
Ann. Bot., May 1, 2006; 97(5): 875 - 881.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
T. Nakano, K. Suzuki, T. Fujimura, and H. Shinshi
Genome-Wide Analysis of the ERF Gene Family in Arabidopsis and Rice
Plant Physiology, February 1, 2006; 140(2): 411 - 432.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
I. A. N. Tonaco, J. W. Borst, S. C. de Vries, G. C. Angenent, and R. G. H. Immink
In vivo imaging of MADS-box transcription factor interactions
J. Exp. Bot., January 1, 2006; 57(1): 33 - 42.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
R. B. Deal, M. K. Kandasamy, E. C. McKinney, and R. B. Meagher
The Nuclear Actin-Related Protein ARP6 Is a Pleiotropic Developmental Regulator Required for the Maintenance of FLOWERING LOCUS C Expression and Repression of Flowering in Arabidopsis
PLANT CELL, October 1, 2005; 17(10): 2633 - 2646.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
T. Nawy, J.-Y. Lee, J. Colinas, J. Y. Wang, S. C. Thongrod, J. E. Malamy, K. Birnbaum, and P. N. Benfey
Transcriptional Profile of the Arabidopsis Root Quiescent Center
PLANT CELL, July 1, 2005; 17(7): 1908 - 1925.
[Abstract] [Full Text] [PDF]


Home page
Plant Cell PhysiolHome page
W.-C. Tsai, P.-F. Lee, H.-I. Chen, Y.-Y. Hsiao, W.-J. Wei, Z.-J. Pan, M.-H. Chuang, C.-S. Kuoh, W.-H. Chen, and H.-H. Chen
PeMADS6, a GLOBOSA/PISTILLATA-like Gene in Phalaenopsis equestris Involved in Petaloid Formation, and Correlated with Flower Longevity and Ovary Development
Plant Cell Physiol., July 1, 2005; 46(7): 1125 - 1139.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
C. Pina, F. Pinto, J. A. Feijo, and J. D. Becker
Gene Family Analysis of the Arabidopsis Pollen Transcriptome Reveals Biological Implications for Cell Growth, Division Control, and Gene Expression Regulation
Plant Physiology, June 1, 2005; 138(2): 744 - 756.
[Abstract] [Full Text] [PDF]


Home page
BioinformaticsHome page
A. Guo, K. He, D. Liu, S. Bai, X. Gu, L. Wei, and J. Luo
DATF: a database of Arabidopsis transcription factors
Bioinformatics, May 15, 2005; 21(10): 2568 - 2569.
[Abstract] [Full Text] [PDF]


Home page
Nucleic Acids ResHome page
S. Nikolajewa, A. Beyer, M. Friedel, J. Hollunder, and T. Wilhelm
Common patterns in type II restriction enzyme binding sites
Nucleic Acids Res., May 11, 2005; 33(8): 2726 - 2733.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
S. de Folter, R. G.H. Immink, M. Kieffer, L. Parenicova, S. R. Henz, D. Weigel, M. Busscher, M. Kooiker, L. Colombo, M. M. Kater, et al.
Comprehensive Interaction Map of the Arabidopsis MADS Box Transcription Factors
PLANT CELL, May 1, 2005; 17(5): 1424 - 1433.
[Abstract] [Full Text] [PDF]


Home page
GeneticsHome page
L. M. Zahn, H. Kong, J. H. Leebens-Mack, S. Kim, P. S. Soltis, L. L. Landherr, D. E. Soltis, C. W. dePamphilis, and H. Ma
The Evolution of the SEPALLATA Subfamily of MADS-Box Genes: A Preangiosperm Origin With Multiple Duplications Throughout Angiosperm History
Genetics, April 1, 2005; 169(4): 2209 - 2223.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
V. Hecht, F. Foucher, C. Ferrandiz, R. Macknight, C. Navarro, J. Morin, M. E. Vardy, N. Ellis, J. P. Beltran, C. Rameau, et al.
Conservation of Arabidopsis Flowering Genes in Model Legumes
Plant Physiology, April 1, 2005; 137(4): 1420 - 1434.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
M. Kooiker, C. A. Airoldi, A. Losa, P. S. Manzotti, L. Finzi, M. M. Kater, and L. Colombo
BASIC PENTACYSTEINE1, a GA Binding Protein That Induces Conformational Changes in the Regulatory Region of the Homeotic Arabidopsis Gene SEEDSTICK
PLANT CELL, March 1, 2005; 17(3): 722 - 729.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
Y. Okushima, P. J. Overvoorde, K. Arima, J. M. Alonso, A. Chan, C. Chang, J. R. Ecker, B. Hughes, A. Lui, D. Nguyen, et al.
Functional Genomic Analysis of the AUXIN RESPONSE FACTOR Gene Family Members in Arabidopsis thaliana: Unique and Overlapping Functions of ARF7 and ARF19
PLANT CELL, February 1, 2005; 17(2): 444 - 463.
[Abstract] [Full Text] [PDF]


Home page
DevelopmentHome page
M. Perez-Rodriguez, F. W. Jaffe, E. Butelli, B. J. Glover, and C. Martin
Development of three different cell types is associated with the activity of a specific MYB transcription factor in the ventral petal of Antirrhinum majus flowers
Development, January 15, 2005; 132(2): 359 - 370.
[Abstract] [Full Text] [PDF]


Home page
DNA ResHome page
K. Iida, M. Seki, T. Sakurai, M. Satou, K. Akiyama, T. Toyoda, A. Konagaya, and K. Shinozaki
RARTF: Database and Tools for Complete Sets of Arabidopsis Transcription Factors.
DNA Res, January 1, 2005; 12(4): 247 - 256.
[Abstract] [Full Text] [PDF]


Home page
J Exp BotHome page
R. Moyle, D. J. Fairbairn, J. Ripi, M. Crowe, and J. R. Botella
Developing pineapple fruit has a small transcriptome dominated by metallothionein
J. Exp. Bot., January 1, 2005; 56(409): 101 - 112.
[Abstract] [Full Text] [PDF]


Home page
Mol Biol EvolHome page
R. C. Moore, S. R. Grant, and M. D. Purugganan
Molecular Population Genetics of Redundant Floral-Regulatory Genes in Arabidopsis thaliana
Mol. Biol. Evol., January 1, 2005; 22(1): 91 - 103.
[Abstract] [Full Text] [PDF]


Home page
Am. J. Bot.Home page
S. Kim, M.-J. Yoo, V. A. Albert, J. S. Farris, P. S. Soltis, and D. E. Soltis
Phylogeny and diversification of B-function MADS-box genes in angiosperms: evolutionary and functional implications of a 260-million-year-old duplication
Am. J. Botany, December 1, 2004; 91(12): 2102 - 2118.
[Abstract] [Full Text] [PDF]


Home page
Genes Dev.Home page
Y. He, M. R. Doyle, and R. M. Amasino
PAF1-complex-mediated histone methylation of FLOWERING LOCUS C chromatin is required for the vernalization-responsive, winter-annual habit in Arabidopsis
Genes & Dev., November 15, 2004; 18(22): 2774 - 2784.
[Abstract] [Full Text] [PDF]


Home page
Am. J. Bot.Home page
W. E. Friedman, R. C. Moore, and M. D. Purugganan
The evolution of plant development
Am. J. Botany, October 1, 2004; 91(10): 1726 - 1741.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
G. Haberer, T. Hindemitt, B. C. Meyers, and K. F.X. Mayer
Transcriptional Similarities, Dissimilarities, and Conservation of cis-Elements in Duplicated Genes of Arabidopsis
Plant Physiology, October 1, 2004; 136(2): 3009 - 3022.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
P. Zimmermann, M. Hirsch-Hoffmann, L. Hennig, and W. Gruissem
GENEVESTIGATOR. Arabidopsis Microarray Database and Analysis Toolbox
Plant Physiology, September 1, 2004; 136(1): 2621 - 2632.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
L. Hennig, W. Gruissem, U. Grossniklaus, and C. Kohler
Transcriptional Programs of Early Reproductive Stages in Arabidopsis
Plant Physiology, July 1, 2004; 135(3): 1765 - 1775.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
V. F. Irish and P. N. Benfey
Beyond Arabidopsis. Translational Biology Meets Evolutionary Developmental Biology
Plant Physiology, June 1, 2004; 135(2): 611 - 614.
[Abstract] [Full Text] [PDF]


Home page
Plant Physiol.Home page
W. Gong, Y.-P. Shen, L.-G. Ma, Y. Pan, Y.-L. Du, D.-H. Wang, J.-Y. Yang, L.-D. Hu, X.-F. Liu, C.-X. Dong, et al.
Genome-Wide ORFeome Cloning and Analysis of Arabidopsis Transcription Factor Genes
Plant Physiology, June 1, 2004; 135(2): 773 - 782.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
S. Ferrario, J. Busscher, J. Franken, T. Gerats, M. Vandenbussche, G. C. Angenent, and R. G.H. Immink
Ectopic Expression of the Petunia MADS Box Gene UNSHAVEN Accelerates Flowering and Confers Leaf-Like Characteristics to Floral Organs in a Dominant-Negative Manner
PLANT CELL, June 1, 2004; 16(6): 1490 - 1505.
[Abstract] [Full Text] [PDF]


Home page
Plant CellHome page
T. Jack
Molecular and Genetic Mechanisms of Floral Control
PLANT CELL, June 1, 2004; 16(suppl_1): S1 - S17.
[Full Text] [PDF]


Home page
Plant Physiol.Home page
A. V. Shchennikova, O. A. Shulga, R. Immink, K. G. Skryabin, and G. C. Angenent
Identification and Characterization of Four Chrysanthemum MADS-Box Genes, Belonging to the APETALA1/FRUITFULL and SEPALLATA3 Subfamilies
Plant Physiology, April 1, 2004; 134(4): 1632 - 1641.
[Abstract] [Full Text] [PDF]


Home page
Proc. Natl. Acad. Sci. USAHome page
J. Nam, J. Kim, S. Lee, G. An, H. Ma, and M. Nei
Type I MADS-box genes have experienced faster birth-and-death evolution than type II MADS-box genes in angiosperms
PNAS, February 17, 2004; 101(7): 1910 - 1915.
[Abstract] [Full Text] [PDF]


Home page
DevelopmentHome page
I. Weir, J. Lu, H. Cook, B. Causier, Z. Schwarz-Sommer, and B. Davies
CUPULIFORMIS establishes lateral organ boundaries in Antirrhinum
Development, February 15, 2004; 131(4): 915 - 922.
[Abstract] [Full Text] [PDF]


Home page
Plant Cell PhysiolHome page
S. Lee, J. Kim, J.-S. Son, J. Nam, D.-H. Jeong, K. Lee, S. Jang, J. Yoo, J. Lee, D.-Y. Lee, et al.
Systematic Reverse Genetic Screening of T-DNA Tagged Genes in Rice for Functional Genomic Analyses: MADS-box Genes as a Test Case
Plant Cell Physiol., December 15, 2003; 44(12): 1403 - 1411.
[Abstract] [Full Text] [PDF]
</