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Europäische Geselischaft für Züchtungsfoschung


Association Européenne pour I’Amélioration des Plantes

PROCEEDINGS OF

CUCURBITACEAE 2016

The XIth Eucarpia Meeting on Cucurbit Genetics & Breeding

Elżbieta U. Kozik & Harry S. Paris


Editors

July 24–28, 2016

Warsaw, Poland
Organizing Committee
Elżbieta U. Kozik, Chair Research Institute of Horticulture
Urszula Kłosińska 1/3 Konstytucji 3 Maja Street
Marcin Nowicki 96-100 Skierniewice, Poland
Marzena Nowakowska www.inhort.pl/en/cucurbit2016
Wojciech Szczechura

Scientific Advisory Committee


Yuling Bai, Wageningen University, The Netherlands
Grzegorz Bartoszewski, Warsaw University of Life Science, Warsaw, Poland
Michael J. Havey, USDA-ARS, University of Wisconsin, Madison, USA
Ales Lebeda, Palacky University in Olomouc, Czech Republic
Amnon Levi, USDA-ARS, Charleston, USA
Stefan Malepszy, Warsaw University of Life Science, Warsaw, Poland
Jordi Garcia Mas, IRTA, Barcelona, Spain
James McCreight, USDA-ARS, Salinas, USA
Katarzyna Niemirowicz-Szczytt, Warsaw University of Life Science, Warsaw, Poland
Harry S. Paris, ARO-Newe Ya’ar, Ramat Yishay, Israel
Zbigniew Przybecki, Warsaw University of Life Science, Warsaw, Poland
Nebehat Sari, Cukurova University, Turkey
Jos Suelmann, Nunhems Seeds/Bayer CropScience, The Netherlands
Yaakov Tadmor, ARO-Newe Ya’ar, Ramat Yishay, Israel
Robert N. Trigiano, University of Tennessee, Knoxville, USA
Todd C. Wehner, North ­­­Carolina State University, USA
Linda Wessel-Beaver, University of Puerto Rico, Mayaguez, PR
Xingping Zhang, Mingkanghui Ecological Agriculture Group, China

Recommended citation style:


In Cucurbitaceae 2016, XIth EUCARPIA Meeting on Genetics and Breeding of Cucurbitaceae,
(Kozik EU and Paris HS, eds), Warsaw, Poland, (pages).
If a figure caption references color(s), please contact the corresponding author for the color version
of the figure.

Printed by:
Wydawnictwo SIGMA Sp. J., Skierniewice, Poland, www.sigma-wydawnictwo.com.pl

ISBN 978-83-7987-896-3
PREFACE

Dear Colleagues,

This book of proceedings contains the full-length contributed papers of the lectures and posters presented
at Cucurbitaceae 2016, the XIth Eucarpia Meeting on Cucurbit Genetics and Breeding. The date of issue of this
book is July 24, 2016, the first day of the conference, following the cucurbit-meeting tradition initiated at the
6th Eucarpia Meeting on Cucurbit Genetics and Breeding (Spain, 1996).
We have edited all of the original contributed papers and contacted all authors for their final approval of the
editing. If, nonetheless, some errors have been overlooked, we apologize.
After the meeting, all the communications printed in these proceedings will be available in PDF format on
the following web site: www.inhort.pl/en/cucurbit2016
The format was considered by us carefully before the distribution of the call for papers. We selected this
format because it is both compact and easily comprehended. We thank all contributing authors for their coop-
eration in preparing their manuscripts according to the format. We acknowledge with pleasure the attention
and professional workmanship of the publisher, Wydawnictwo Sigma, in helping us produce what we believe
to be a fine book of proceedings.

July 4, 2016

Editors
Elżbieta U. Kozik
Harry S. Paris
Eucarpia Cucurbitaceae 2016 Conference
was organized under the auspices of:

Vegetable Section of Research Institute


EUCARPIA City of Horticulture
of Skierniewice

We are grateful to our sponsors for their generous


and ongoing support of Cucurbitaceae 2016

GOLD SPONSORS

SILVER SPONSOR

CONFERENCE PARTNERS
ORGANIZING COMMITTEE MESSAGE

Dear Colleagues,

We, the Organizing Committee of Cucurbitaceae 2016, the XIth Eucarpia Meeting on Cucurbit Genetics and
Breeding, welcome you back to Warsaw, Software crack - Activators Patch, where the Windows 8.1 crack - Free Activators such meeting was held in 1992.
The Eucarpia meetings on Cucurbit Genetics and Breeding were initiated in the 1970s. Since 1988, they
have been held on a regular basis every four years, in France (1988), Poland (1992), Spain (1996), Israel
(2000), Czech Republic (2004), France (2008), Turkey (2012), and now again Poland. Meetings on cucurbit
genetics and breeding have been held in the United States of America since the 1980s. Since 1994, they have
been held on a regular basis in the even-numbered years alternating with the Eucarpia meetings, in Texas
(1994), California (1998), Florida (2002), North Carolina (2006), South Carolina (2010), and Michigan (2014).
We greatly appreciate the financial support of the EUCARPIA Cucurbitaceae 2016 Sponsors. Please join
us expressing thanks to the Sponsors` representatives for their generous contributions.
As of this writing, the following have furnished much-needed financial and material support for
Cucurbitaceae 2016: Eucarpia, InHort Skierniewice, City of Skierniewice, Bayer, Amy Goldman Fowler,
Bejo, Syngenta, Rijk Zwaan, Polan Kraków, PNOS, PlantiCo, SANLAB, RB, and TORSEED.
We hope Cucurbitaceae 2016 will be the most memorable and enjoyable conference for all participants.

July 4, 2016
Organizing Committee
Elżbieta U. Kozik
Urszula Kłosińska
Marcin Nowicki
Marzena Nowakowska
Wojciech Szczechura
Table of Contents

ORAL PRESENTATIONS cmv1 Encodes a Vacuolar Protein Sorting 41 Involved


in Transport of Cucumber Mosaic Virus in Melon. 54
Session I: History and Breeding Ana Giner, Laura Pascual, Gabor Gyetvai, Michael
Bourgeois, Jordi Garcia-Mas, and Ana Montserrat
History of Cucumber Breeding in Poland. 11
Martin-Hernández
Elżbieta U. Kozik
Emergence of the Sweet Dessert Watermelon, Session IV: Breeding and Genetic Resources 3
Citrullus lanatus, in Mediterranean Lands During
the Roman Era.16 Genome Sequence of the Bottle Gourd
Harry S. Paris [Lagenaria siceraria (Mol.) Standl.] and Applications
to the Study of Hetero-Graft-Conferred Cold Tolerance.58
Status of Cucurbit Breeding at AVRDC – The World Pei Xu, Nailin Xin, Xiaohua Wu, Xinyi Wu, Ye Tao,
Vegetable Center.21 Yunping Huang, Baogen Wang, Yaowen Hu,
Narinder P.S. Dhillon, Supannika Sanguansil, Zhongfu Lu, Yuhong Wang, and Guojing Li
Supornpun Srimat, Hsin-chun Cheng, Chung-cheng Lin,
Ramasamy Srinivasan, Lawrence Kenyon, New Sources of Resistance to CYSDV in Melon. 61
Roland Schafleitner, Ray-yu Yang, and Peter Hanson James D. McCreight, William M. Wintermantel,
and Eric T. Natwick
Session II: Breeding and Genetic Resources 1 Salt Stress Affects the Chloroplast Ultrastructure
of Different Ploidy Watermelon Leaves.66
Superfruiter (sf): A New Melon Type with Multiple
Hong-ju Zhu, Wen-ge Liu, Sheng-jie Zhao, Zhi-hong
Small Seedless Fruits and Increased Yield.26
Yan, Xu-qiang Lu, Nan He, and Ping-li Yuan
Ya’akov Tadmor, Ido Oz, Uzi Sa’ar, Ayala Meir,
Fabian Baumkoler, Nurit Katzir, Efraim Lewinsohn,
Vitaly Portnoy, Galil Tzuri, Arthur A. Schaffer, Session V: Genomics 1
and Joseph Burger Progress in Assembling the Cucumber (Cucumis sativus)
The Andromonoecious Allele of CitACS4 Reduces Seed Borszczagowski B10 Line Genome Using Long Single
and Fruit Set in Watermelon.29 Molecule, Real-Time Reads.72
Encarnación Aguado, Susana Manzano, Alicia García, Paweł Osipowski, Michał Wojcieszek, Magdalena
and Manuel Jamilena idm 6.33 crack - Free Activators Pawełkowicz, Agnieszka Skarzyńska, Sergey Koren,
Alex Lomsadze, Rafał Wóycicki, Wojciech Pląder,
The Rhizosphere and Seed Microbiomes of Oil Pumpkin Kohei Yagi, Mark Borodovsky, Stefan Malepszy,
Breeding Lines, Hybrids, and a Population Cultivar. 34 and Zbigniew Przybecki
Eveline Adam, Maria Bernhart, Henry Müller,
Johanna Winkler, and Gabriele Berg Progress in Cucumber Molecular Breeding.75
Xingfang Gu, Shengping Zhang, Han Miao, and Ye Wang
Inheritance of Rind Pattern in Watermelon.38
Todd C. Wehner and Lingli Lou The Plant Breeders’ Dream Come True: Editing
Non-Transgenic Cucumber for Broad Virus Resistance
Significant Reciprocal-Cross Differences in Cucumber.43 Using CRISPR/Cas9 Technology.80
Jia Shen, Rob Dirks, and Michael J. Havey driverdoc activation key free - Free Activators Dalia Wolf, Jeyabharathy Chandrasekaran,
Marina Brumin, Diana Leibman, Chen Klap,
Session III: Breeding and Genetic Resources 2 Mali Pearlsman, and Amit Gal-On
Isolation and Characterization of Three Recessive QTL Mapping of Cucumber Parthenocarpy
Andromonoecious Mutants of Cucurbita pepo.45 by QTL-Seq.86
Alicia García, Susana Manzano, Encarnación Aguado, Min He, Qianya Liu, Xiaohua Qi, Qiang Xu,
Zoraida Megías, Cecilia Martínez, Dolores Garrido, and Xuehao Chen
and Manuel Jamilena
Genome-Wide Association Study of Powdery Mildew Session VI: Genomics 2
Resistance in a Worldwide Collection of Melon A Chromosome Translocation Leads to Gynoecy
(Cucumis melo L.) Germplasm.50 in Watermelon.89
Qianglong Zhu, Chuan Wu, Sikandar Amanullah, Jie Zhang, Gaojie Ji, Hong Zhao, Honghe Sun,
Shi Liu, Peng Gao, Xuezheng Wang, Hongyan Ma, Shouwei Tian, Guoyi Gong, Jianting Shi, Shaogui Guo,
Zicheng Zhu, and Feishi Luan Yi Ren, Haiying Zhang, and Yong Xu
S-Gene and R-Gene Candidates for Disease Sensitivity to Fungicides in Podosphaera xanthii
Resistance in Watermelon.94 and Impact on Managing Cucurbit Powdery Mildew
Henk J. Schouten, Jeroen A. Berg, and Yuling Bai in New York, USA. 144
Margaret T. McGrath
Dissection of Climacteric and Non-Climacteric
Ripening in Melon by Using Genetic, Analytic, Initiative for International Cooperation of Researchers
and Genomic Resources.100 and Breeders Related to Determination
Lara Pereira, Sergi Pérez, Pablo Rios, and Denomination of Cucurbit Powdery Mildew
Konstantinos Alexiou, Marta Pujol, Michael A. Phillips, Races.148
and Jordi Garcia-Mas Aleš Lebeda, Eva Křístková, Božena Sedláková,
and James D. McCreight
Utilizing Genetic Diversity in the Desert Watermelon
Citrullus colocynthis for Enhancing Watermelon Survival of Xanthomonas cucurbitae, Cause
Cultivars for Resistance to Biotic and Abiotic Stress.105 of Bacterial Spot of Cucurbits, in the Field.153
Amnon Levi, Alvin Simmons, Kai-shu Ling, Ya’akov Sita Thapa and Mohammad Babadoost
Tadmor, Padma Nimmakayala, and Umesh K. Reddy
Inheritance and QTL Mapping of Resistance Session IX: Biotic Stresses 3
to Gummy Stem Blight in Cucumber
Molecular Characterization of the P1 Protein of
(Cucumis sativus L.).109
the Zucchini Yellow Mosaic Virus in Turkey. 157
Shengping Zhang, Shulin Liu, Han Miao, Min Wang,
Serife Topkaya and Filiz Ertunc
Yanxia Shi, Ye Wang, Baoju Li, and Xingfang Gu
Sensitivity of Phytophthora capsici Isolates
from Illinois to Fungicides.161
Sessions VII: Biotic Stresses 1
Jose R. de Souza, Tiyoko N.H. Rebouças,
Global Occurrence of the A2 Mating Type Yiwen Xiang, and Mohammad Babadoost
of Pseudoperonospora cubensis, the Causal Agent
of Cucurbit Downy Mildew. 115 A Point Mutation of the Coat Protein of Cucumber
Green Mottle Mosaic Virus Alters the Symptoms
Yigal Cohen, Avia E. Rubin, Lidan Falach,
from Mosaic to Yellow Spots. 166
Mariana Galperin, and Fabian Runge
Liming Liu, Bin Peng, and Qinsheng Gu
Genetic Structure of Pseudoperonospora cubensis
Populations Infecting Commercial Sources of Variation in ELISA Tests Used to Quantify
and Non-Commercial Hosts in North Carolina. 119 trojan remover free download full version with key - Free Activators ZYMV and PRSV Resistance in Cucurbita moschata.170
Emma C. Wallace and Lina M. Quesada-Ocampo Linda Wessel-Beaver and Jose Carlos V. Rodrigues

Interactions of Cucumis melo and Cucurbita


spp. Accessions with Pseudoperonospora cubensis Session X: Quality Fruit
Are Race- (Pathotype-) Specific.123 Metabolomic Plasticity of Cucumber Fruit Peel -
Aleš Lebeda, Eva Křístková, Jana Štěpánková, Effects of Developmental Stage and Market Class. 174
Božena Sedláková, Jana Roháčková, Harry S. Paris, Ben N. Mansfeld and Rebecca Grumet
and Mark P. Widrlechner
Towards Whole-Genome Association Mapping
Cellular and Biochemical Mechanisms of Multiple Fruit Quality Traits in Melon and Squash.180
of Cucumber Resistance Against Amit Gur, Harry S. Paris, Galil Tzuri, Ayala Meir,
Pseudoperonospora cubensis .128 Nurit Katzir, Vitaly Portnoy, Efraim Lewinsohn,
Marcin Nowicki, Urszula Kłosińska, Wojciech Arthur A. Schaffer, Joseph Burger, and Ya’akov Tadmor
Szczechura, Monika Markiewicz, Iwona Sowik,
Lech Michalczuk, and Elżbieta U. Kozik Mapping the Major Genes Related to Lycopene
Content and Flesh Color Traits in Watermelon
(Citrullus lanatus). 186
Sessions VIII: Biotic Stresses 2 Shi Liu, Qianglong Zhu, Wu Chuan,
Loss-of-Function Mutations in a Cucumber MLO Sikandar Amanullah, Peng Gao, Xuezheng Wang,
Gene Lead to Hypocotyl Resistance to Powdery Hongyan Ma, Zicheng Zhu, Feishi Luan,
Mildew .132 and Angela R. Davis
Jeroen A. Berg, Yuling Bai, and Henk J. Schouten Extreme Picture Finder 3.51.4 Crack - Activators Patch Effects of Grafting on Carbohydrate Metabolism
Expression Analysis on Melon MLO Family Genes in Melon (Cucumis melo L.) During Stages of Fruit
and Identification of CmMLO2 Mutants Resistant Development.191
to Powdery Mildew .138 Qiushi Fu, Xinying Zhang, Qiusheng Kong, Zhilong Bie,
Hong Cheng, Wei-ping Kong, Hong-zhong Yue, and Huaisong Wang
and Jun-Feng Lü
POSTER PRESENTATIONS Searching for Candidate Genes Controlling Flower
and Fruit Development in Cucurbita pepo.237
Effective Long-Term Storage Methods for Pollen Cecilia Martinez, Cristina Esteras, Susana Manzano,
in Bitter Gourd (Momordica charantia L.).197 Manuel Jamilena, and María Belén Pico
Masako Akutsu Identification and Mapping of a QTL in Cucumber
Notes on in vivo and in vitro Tetraploidization of Bottle Controlling Resistance to Pseudoperonospora cubensis . 242
Gourd (Lagenaria siceraria), Citron Watermelon Wojciech Szczechura, Urszula Kłosińska,
(Citrullus lanatus var. citroides), and Colocynth Marzena Nowakowska, and Elżbieta U. Kozik
(C. colocynthis).199
Effects of Some Plant Activators on Yield, Plant,
Mohamed Dhamir Kombo, Nebahat Sari, Baris Dal, and Fruit Characteristics of Summer
and Ilknur Solmaz and Winter Melons. 245
Double-Haploid Pure-Line Development in Galia-Type Ceren Ayse Bayram and Nebahat Sari
Melons Having a Long Shelf Life.205
Effects of Cadmium Stress on Growth, Reactive
Seher Pamuk, Nebahat Sari, Ibrahim Mancak, Oxygen Metabolism, and Photosynthesis
Ilknur Solmaz, Baris Dal, and Ahmad Alsaleh of Melon Seedlings. 250
Genetic Diversity Analysis of Winter Squash Yongping Zhang, Youyuan Chen, and Weihong Gu
(Cucurbita maxima Duchesne) Accessions Using SSR
Construction of a High-Density DArT-seq
Markers.210
SNP-Based Genetic Map in a Population Derived
Karolina Kaźmińska, Krzysztof Sobieszek, from a Cross Between a Feral and a Cultivated
Małgorzata Targońska, Aleksandra Korzeniewska, Watermelon.255
Katarzyna Niemirowicz-Szczytt,
Runsheng Ren, Rumiana Ray, Pingfang Li, Jinhua Xu,
and Grzegorz Bartoszewski
Man Zhang, Guang Liu, Xiefeng Yao, Andrzej Kilian,
Inheritance of Tolerance to Tomato Leaf Curl New Delhi and XingpingYang
Virus (ToLCNDV) in Melon. 214
Effects of Low Night Temperature on Morphological
Cristina Sáez, María Ferriol, Cecilia Martínez, Anatomy Structure of Grafted Watermelon
Cristina Esteras, Carmelo López, and María Belén Picó Seedlings.261
Differential Effects of Drought Stress on Germination Xingping Yang, Xuedan Hu, Man Zhang, Jinhua Xu,
and Seedling Growth of Cucumber Accessions. 217 Runsheng Ren, Guang Liu, Qian Hou, Xiefeng Yao,
Urszula Kłosińska, Elżbieta U. Kozik, Waldemar Treder, Pingfang Li, and Xuehao Chen
and Krzysztof Klamkowski Analysis of Fruit-Flesh Aroma Profile in a Melon
CucCAP - Developing Genomic Resources Core Collection .265
for the Cucurbit Community. 222 Cristina Vso downloader 5.0.1.53 crack - Crack Key For U, José Luis Rambla, Gerardo Sanchez,
Rebecca Grumet, Zhangjun Fei, Amnon Levi, Antonio Granell, and María Belén Picó
James D. McCreight, Michael Mazourek, Marco Palma, Long-Lasting (2001 to 2009) Variation in Virulence
Jonathan Schultheis, Yiqun Weng, Mary Hausbeck, Among Czech Cucurbit Powdery Mildew Populations
Shaker Kousik, Kai-Shu Ling, Cecilia McGregor, Screened on Eleven Cucumis melo Differential
Lina M. Quesada-Ocampo, Angela Linares Ramirez, Genotypes . 268
Umesh K. Reddy, Louis Ribera, Christine Smart,
Božena Sedláková, Eva Rušáková, Eva Křístková,
Pat Wechter, Todd C. Wehner, Linda Wessel-Beaver,
and Aleš Lebeda
and William M. Wintermantel
Comparison of Anatomic Structure and Enzyme Activity
Germination of Wild Cucumis Species
of Fruit Rinds in Crack-Resistant and Crack-Prone
and Interspecific Hybrids.227
Watermelons.272
Andrés Cáceres, María Belén Pico, and Carmina Gisbert
Meiling Gao, Changbao Yu, Xiaoming Wei, Jiayi Li,
Evaluation of a Cucumber RILs Population for Yan Liu, and Yu Guo
Resistance to Angular Leaf Spot. 231
Determination of Salt Tolerance Potential of Turkish
Renata Słomnicka, Helena Olczak-Woltman, Aleksandra Bottle Gourd (Lagenaria siceraria) Germplasm .278
Korzeniewska, Katarzyna Niemirowicz-Szczytt,
Halit Yetişir, Nihal Denli, Atilla Ata, Abdullah Ulaş,
and Grzegorz Bartoszewski
Hasan Pinar, Halil Kirnak, and Mehmet Yamaç
Utilization of a Strongly Female Mutant
Open-Field Survey of Turkish Bottle Gourd
in Seed-Pumpkin Germplasm of Cucurbita maxima
Germplasm Reaction to Virus Diseases. 283
Duchesne for Increased Production and to Facilitate
Hybrid Seed Production. 234 Hakan Fidan, Nihal Denli, Halit Yetisir, Pelin K. Ozturk,
and Çetin Nacar
Guoyu Zhang, Haizhen Li, Fan Zhang, and Jing Wang
Reaction of Some Melon (Cucumis melo) Genotypes Effect of Melon Rootstocks with Multiple Disease
to Drought Stress.288 Resistance on Yield and Quality of Melon
Muhemet Zeki Karipcin, Nebahat Sari, (Cucumis melo L.). 322
and Ilknur Solmaz Su Kim, Ji Hye Moon, Yun Chan Huh, Yoon Ah Jang,
Se Woong An, Dong Kum Park, Tae Sung Park, Eun
Evaluation of Genetic Relationships Among Hungarian
Young Yang, Myeong Cheoul Cho, and Soo Young Chae
and Turkish Watermelon (Citrullus lanatus var. lanatus)
Accessions by SSR Markers.292 Breeding of F1 Hybrid Melon ‘Greenpower’ with
Csaba Szamosi, Ozhan Simsek, Yildiz Aka Kacar, Multiple Disease Resistance as Rootstock for Melon
Ilknur Solmaz, and Nebahat Sari (Cucumis melo L.).325
Yun Chan Huh, Su Kim, Ji Hye Moon, Yoon Ah Jang,
Genome-Wide Association Mapping of Free Glutamic
Dong Kum Park, Dae Young Kim, Eun Young Yang,
Acid Content Responsible for Umami Taste in Bottle
Hee Joo Lee, and Won Byoung Chae
Gourd.297
Xinyi Wu, Pei Xu, Xiaohua Wu, Baogen Wang, Comparison of Some Cucumber Cultivars for
Zhongfu Lu, and Guojing Li Open-Field Production Present on the Polish
National List. 331
Use of Wild Cucumis as Potential New Rootstocks
Katarzyna Bartoszak
for Melons. 300
Andrés Cáceres, María Ferriol, Carmina Gisbert, Detection of Two QTLs Associated with Resistance
and María Belén Pico to Cucurbit yellow stunting disorder virus in Melon
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Evaluation of Organically Produced Melon Cultivars
Francisco J. Palomares-Ríus, Ana Garcés-Claver,
for Powdery and Downy Mildew Severity in Maryland,
and Maria L. Gómez-Guillamón
U.S.A.305
Sasha C. Marine and Kathryne L. Everts Mutation in a Conserved Position
of 1-Aminocyclopropane-1-Carboxylate Synthase
Association Analysis of Morphological Traits Leads to Andromonoecy in Watermelon.338
in Iranian Extreme Picture Finder 3.51.4 Crack - Activators Patch Accessions. 308
Gaojie Ji, Jie Zhang, Haiying Zhang, Guoyi Gong,
Masoud Maleki, Abdolali Shojaeiyan, Extreme Picture Finder 3.51.4 Crack - Activators Patch Shaogui Guo, Yi Ren, and Yong Xu
and Sajad Rashidi-Monfared
Breeding of New Watermelon Hybrid ‘Shenmi-968’
Identification of Informative ISSR Markers for with Disease Resistance and High Quality .342
Morphological Characteristics of Iranian Dudaim
Wei-hong Gu, Rong-hao Song, Hong-juan Yang,
Melons (Cucumis melo var. dudaim).312
Li-hua Zhu, and Chao-han Li
Fateme Mirnouri, Abdolali Shojaeiyan, Masoud Maleki,
and Azam Nikzad Gharehaghaji A New Type of Semi-Bush Habit Processing Drivermax pro serial keygen - Free Activators (Cucumis sativus) for Simplified Fruit Harvest.347
Identification of ISSR Markers Linked to
Aleksandra Korzeniewska, Teresa Gałecka, Ewa
Morphological Traits in Iranian Snake Melons.318
Guzowska, and Katarzyna Niemirowicz-Szczytt
Winzip 21.5 registration code - Crack Key For U Dastranji, Abdolali Shojaeiyan,
and Mohsen Falahati-Anbaran
AUTHOR INDEX .351
History of Cucumber Breeding in Poland
winzip 21.5 registration code - Crack Key For U Elżbieta U. Kozik
Research Institute of Horticulture, 1/3 Konstytucji 3 Maja, 96-100 Skierniewice, Poland
e-mail: elzbieta.kozik@inhort.pl

Abstract. Cucumber (Cucumis sativus) is the most economically important cucurbit in Poland and has been
cultivated in this country for centuries. Since the 19th century, companies and growers alike maintained and
reproduced seeds of vegetable cultivars of local and foreign origin, mainly from Denmark, Holland, and Germany.
Polish cucumber breeding was initiated only at the beginning of the 20th century, by three private companies,
“C. Ulrich”, “Hoser Brothers”, and “E. Freege”. Just before World War II (1939), the first Polish register of original
vegetable varieties was established; it included both first Polish field cucumbers ‘Przybyszewski Hoser’ (“Hoser
Brothers”) and ‘Warszawski’ (“C. Ulrich”). But, the history of cucumber breeding in Poland resembles the history
of our country, turbulent and demanding sacrifices. Unfortunately, in the 1950s, all private breeding companies
were nationalized. The successors of the three original privately owned Polish breeding companies underwent
numerous administrative transformations, becoming State Treasury-owned commercial enterprises before the
new millennium, and they operate in this form today. Herewith presented are the achievements of individual
Polish companies and academic institutions in breeding improved cultivars for the cucumber production industry.
Keywords: Cucumis sativus, cultivars, F1 hybrids, genetic resources, processing industry

Introduction growers in Poland imported seeds, including cucumber, mainly


for their own production and reproduction as they saw fit.
Cucumber (Cucumis sativus L.) is the most important cu- Some knowledge of the advanced level of gardening in western
curbit in Poland, being commonly cultivated in large commer- Europe did reach Poland, invoking considerable interest among
cial operations as well as home gardens, for processing and gardening enthusiasts. This spurred increasingly frequent trips
fresh market. According to FAOSTAT (www.faostat.fao.org) to accrue knowledge and skills that would benefit local horticul-
statistics for 2012, Poland was first in the European Union for ture. The private seed companies “C. Ulrich”, “Hoser Brothers”,
area planted with cucumber (17,100 ha) and second for produc- and “E. Freege”, in the 19th century, were the first establishments
tion (520,868 t). Over the past 10 years, the area planted with to promote the introduction and adoption of horticultural plants
cucumber has slightly decreased, while productivity slightly in- in Poland. Their present-day successors continue this important
creased. Annual consumption of cucumbers in Poland is about function.
7 kg/person. “C.  Ulrich”, founded in 1805 in Warsaw, was the first Pol-
ish seed company. Initially, the company specialized in vegeta-
ble produce, fruits, and ornamental plants, but starting in 1830
Cucumber adoption and the beginnings it introduced the sale of vegetable seeds imported from abroad,
of cucumber breeding in Poland mainly from Germany, Denmark, and the Netherlands. The com-
pany launched actual breeding activities of vegetables, including
Cucumbers probably first entered Poland by the overland cucumber, in the 1930s (Korohoda 1984).
route across southwestern Asia through to eastern Europe in “Hoser Brothers” was established in Warsaw in 1848. Early
early medieval times (Paris et al. 2012). Cucumbers have been on, it engaged in the production of vegetable produce and orna-
grown in Poland since then. Maintenance and seed reproduction mental plants. The beginning of the 20th emsisoft anti-malware 2019 crack - Activators Patch saw expansion
of cucumbers and other vegetables occurred later in Poland than to its actual breeding of vegetable and ornamental plants, and
in some other European countries, due to its cool, unfavorable a marked increase in production of its seed varieties.
climate and history of political upheaval. During the 19th centu- “E. Freege” was established in 1860 in Cracow. At first it was
ry, various European companies were established that special- a horticultural farm, shifting its activities over time towards seed
ized in the maintenance and reproduction of old, established production. This meant, in essence, import of the best foreign
vegetable cultivars and breeding new, improved ones. Vegetable varieties and their sale, a model of business similar to English
and German seed companies.
In 1917, in the western part of Poland, the oldest of the
currently operational major vegetable market companies was
E.U. Kozik and H.S. Paris (Eds.): Proceedings of Cucurbitaceae 2016,
the XIth EUCARPIA Meeting on Genetics and Breeding of Cucurbitaceae. founded: “Spójnia Nochowo”. Its main activity then was the
July 24-28, 2016, Warsaw, Poland. production of vegetable seeds. During the inter-war period, this

11
company was the only breeding facility in western Poland, and lished the Research Center for Cultivar Testing (Polish acronym
its seed reproduction activities contributed greatly to raising the COBORU) near Poznań, which operates to this day. Evaluations
status of gardening in this country. of cultivars sent to the Register were carried out at the time in
At the time, plant breeding in the true sense was poorly devel- different parts of Poland. Cucumber cultivars were evaluated in
oped in Poland. Producers of vegetable seeds, including cucum- Puławy, in the east. Comparison standards were cultivars impor-
bers, mainly imported cultivars from abroad and reproduced them ted from Denmark, Germany, and the Netherlands. Based on the
at their own discretion. Then, in 1937, just before the outbreak of results, the first selection of vegetable cultivars was made at the
the World War II, the Polish Association of Seed Producers de- Research Institute of Vegetable Crops (RIVC) in Skierniewice,
cided to carry out an assessment of seed samples obtained from led since its inception in 1964 by Prof. Emil Chroboczek. Later
Polish breeders. Only a small part of these was then selected, on and until now, COBORU publishes annually a list of vegeta-
after thorough research conducted in the Department of Crop and ble varieties registered in Poland. In Poland, as in the other EU
Vegetable Breeding (Warsaw University of Life Sciences, then in countries, The Common Catalog of Varieties of Vegetable Plant
Skierniewice), for the national Preliminary Register of Original Species is in effect.
Varieties, the first Polish register of original vegetable cultivars. Currently, most cucumber breeding in Poland is conducted at
Resulting from this assessment of the seed samples submitted by five centers: three seed companies operating as Treasury-owned
domestic breeders, 35 cultivars of vegetables were accepted into commercial companies (“POLAN Cracow”, “PlantiCo Zielon-
the Register, including two field cucumbers, ‘Hoser Przybysze- ki”, and “Spójnia Nochowo”) and two free audio converter crack - Crack Key For U institutions
wski’ (“Hoser Brothers”) and ‘Warszawski’ (“C. Ulrich”). (Warsaw University of Life Sciences and the Research Institute
Only in the three years 1936–1939 did vegetable cultivation of Horticulture Skierniewice, InHort).
experience intensive development, and it is this period that is
considered the birth of the new era of plant breeding in Poland. “POLAN Cracow”
At this time, many species, including cucumber, began to under- Field cucumber has been and still is the most import-
go dynamic improvement. Despite great difficulties, and owing ant vegetable crop of the company assortment. Its first culti-
to the erstwhile commitment of various groups of farmers and vars, ‘Delicius’, ‘Mikor’, and ‘Monastyrski’, all monoecious,
breeders, Poland managed to almost catch up with the European were registered in the 1970s (Żuradzka 2016). For many years
level of plant breeding. The first export of Polish vegetable seed (1975–2008), ‘Monastyrski’ has been widely grown in Poland,
varieties was planned for the fall of 1939. No one suspected, being used in processing. Introduction of F1 hybrids, much more
though, that the outbreak of WWII would almost completely productive than the traditional monoecious cultivars, was a rev-
destroy these new achievements and impede plant breeding in olution in cucumber cultivation. The first Polish-bred F1 hybrid
Poland for decades. cucumber, ‘Polan’, from the RIVC Skierniewice (currently
InHort) was bred by Prof. Bogdan Kubicki and Dr. Ewelina
Kłossowska. This cultivar was registered in 1972 and licensed
Cucumber breeding after 1945 to “POLAN Cracow”. ‘Polan F1’ is still very popular among
growers. Soon thereafter, Dr. Izabela Żuradzka of “POLAN
World War II hostilities and the Warsaw Uprising (1944) Cracow” led a team of cucumber breeders, in collaboration with
resulted in severe losses, especially for the companies that had Prof. Kubicki, in the development and registration of several
facilities located near Warsaw, but also throughout the country. high-yielding cucumber hybrids of various fruit types, with re-
Only small quantities of breeding materials were rescued. Pol- sistance to Cladosporium cucumerinum (scab): ‘Dragon’ (1976),
ish plant breeding suffered acutely from the loss of outstanding ‘Mieszko’ and ‘Racibor’ (both 1979). Fifteen to 20 years later,
professionals who died at the hands of the occupiers. The surviv- “POLAN Cracow” released seven pickling cucumber F1 hybrids
ing older generation and the young people educated by them had with resistance to Pseudoperonospora cubensis (downy mildew)
to rebuild Polish seed science and plant breeding almost from and Cucumber mosaic virus (CMV): ‘Krak’, ‘Julian’, ‘Borus’,
scratch, in very difficult post-war conditions. Unfortunately, too, ‘Zagłoba’, ‘Hubal’, ‘Chrobry’, and ‘Forum’.
in the 1950s, all private companies were nationalized. Under An indubitable success of the company was the development
state aid, they were offered leases of a number of farms, where of the first Polish gherkin-type F1 hybrids, ‘Anulka’, ‘Rufus’,
they could organize new plant breeding stations. On the basis of and ‘Tytus’. These were very well received by growers due to
“E. Freege”, decades later and after many administrative and or- high yields, general pathogen tolerance, and high fruit quality.
ganizational transformations, “POLAN Cracow” was established Another major commercial achievement was for pickling cu-
in 1992, and is operational today. On the basis of the nationalized cumbers, ‘Julian F1’ (1996) being exceptional because of its very
assets of “Hoser Brothers” and “C.  Ulrich”, nine horticultural good fruit processing quality.
plant breeding stations were created in the 1950s, which even- In the late 1980s, the company started cooperation with the
tually became part of “PlantiCo Zielonki” of Warsaw. “PlantiCo RIVC Skierniewice because of problems with Pseudoperono-
Zielonki” is the only survivor of the former nucleus of Warsaw spora cubensis infestations, which decimated cucumber plant-
plant breeding. As early as January 1945, “Spójnia Nochowo” ings all over Poland. The work focused on methods for screen-
re-undertook its activity, but at the end of 1950 it came under ing for resistance. Efforts were conducted under the direction of
compulsory state control and then nationalized. Many organiza- Prof. Roch W. Doruchowski. After his retirement in 2000, the
tional transformations followed. Ultimately, in 1994, it became a cooperation continued, coordinated until this day by myself.
State Treasury-owned company, functioning in this form today. Common interests cover a search for new sources of resistance
The State Register of Original Varieties was established in to P. cubensis, determining the mode of inheritance of resistance,
Poland in 1952. In 1966, the Polish Ministry of Agriculture estab- identifying DNA markers for resistance, insights into the genet-

12
ic diversity of the pathogen in Poland, and generation of new boom. The hit field pickling cucumber ‘Śremski F1’ (1988) is
genetic variability in cucumber for low-temperature tolerance. the major reason. Popularity of this early variety among pro-
As a result of this cooperation, the company received a wealth cessors lasts to this day, keeping the company in good finan-
of valuable germplasm for their breeding efforts. In 1999, the cial condition. ‘Śremski F1’ has proven to be an outstanding
company undertook cooperation with the Department of Plant cultivar; the seed sales in the years 1998 to 2015 amounted to
Genetics, Biotechnology, and Breeding (Warsaw University of 350 t, representing 24% of the seed sales of field cucumbers in
Life Sciences), to obtain parental lines of double haploids (DH) Poland. Notable progress was accomplished during 1994–2004,
by induced parthenogenesis. Currently, such lines are being with the development and release of three new parthenocarpic and
developed in vitro by the company. Members of the cucumber genetically devoid of bitterness F1 hybrid cucumbers for green-
breeding team of the company include Marta Antos, Eugenia house production: ‘Orion’, ‘Polonez’, and ‘Szafir’. Subsequent-
Koterwa, and Aleksander Kopta. In all, “POLAN Cracow” has ly, the company established its own in vitro laboratory, in which
bred 40 cultivars of field cucumbers, including 37 F1s, of which they fast-track the breeding with DH lines. They also have set up
20 are widely cultivated. a resistance-breeding laboratory, working towards the increase
in the level of resistance to major cucumber pathogens, mainly
“PlantiCo Zielonki” Pseudoperonospora cubensis. The results of this work are four F1
An heir to the tradition of “C. Ulrich”, “PlantiCo Zielonki” hybrids with resistance against P. cubensis, ‘Śremianin’, ‘Lider’,
took over the pre-war registered field cucumber ‘Warszawski’ ‘Husar’, and ‘Meteor’, which were released from 2002 to 2008.
(Karpiński et al. 2016). The company also carried large collections In 2002, “Spójnia” also released the first two Polish parthenocar-
of foreign cultivars, of which ‘Delicius’ (1970) and ‘Wisconsin’ pic cucumbers, ‘Rubin F1’ and ‘Jaspis F1’, for cultivation under
(1973) were registered at COBORU. The 1970s mark the origins low covers and recommended for low-salt souring (fermentation
of breeding of its own cucumber cultivars. During that decade, for only a few days). The low-salt souring group was then joined
“PlantiCo” registered four F1 salad cucumber hybrids: ‘Fryko’, by ‘Lazuryt F1’ (2004), another hybrid having short fruits and for
‘Ikar’, ‘Kometa’, and ‘Kuba’, that were bred in collaboration with cultivation primarily under low tunnels, and ‘Bursztyn F1’ and
the RIVC Skierniewice. In the 1980s and ‘90s, the company mark- ‘Karat F1’ (both in 2007) for open field production. In 2009–2014,
edly increased its cucumber breeding activities and released 24 the company introduced a series of eight new early cucumbers
cucumber cultivars, which makes 53% of all those registered by suitable for pickling, and its latest additions are early, high-yield-
the company. The major economic achievements of the company ing, and tolerant to P. cubensis ‘Atut F1’ and ‘Śremski Nowy F1’.
are its five hybrids for greenhouse production ‘Replika’, ‘Atos’, Overall, the company has registered 42 hybrids, including 27 field
‘Caruso’, ‘Heros’, and ‘Ines’. These bridged the gap in the assort- cucumbers, nine greenhouse type, and six for cultivation under
ment of greenhouse cucumber cultivars and created alternatives plastic tunnels. The largest contributors to the company breeding
to the Dutch semi-long type. Other notable achievements include efforts have been: Marian Pędziński (40 cultivars), Anna Nowak
breeding of new F1 cultivars of field cucumber: smooth skin, no (15 cultivars), Arkadiusz Labrzycki (15 cultivars), Szymon Frąc-
warts, for salad ‘Gracius’ (1992) and for pickling ‘Soplica’ (2001). kowiak (13 cultivars), and Wojciech Matuszak (6 cultivars).
The main contributors to “PlantiCo’s” broad array of cucumber
cultivars are: Barbara Chrzanowska, Jadwiga Tomczyk, Teresa Other companies
Bartłomiejczak, and Maria Guzik. So far, the company has re- Out of the 110 cucumber cultivars currently listed in the
leased 45 cucumber hybrids, including 29 of field cucumbers and COBORU Register (February, 2016), 38 were registered by es-
16 greenhouse cucumbers. tablishments other than the three described above. These include
For many years, “PlantiCo” cooperated with academia in Polish breeding and seed companies, and some private breeding
cucumber breeding. Prof. Katarzyna Niemirowicz-Szczytt (War- enthusiasts. It is important to note that foreign companies may
saw University of Life Sciences) guided the creation of a range also register in the Common Catalog of Varieties of Vegetable
of DH lines with their subsequent takeover by PlantiCo. Dr. Ur- Species, and such registration is obligatory for each European
szula Kłosińska and I at InHort bred cucumbers for increased Union country.
resistance to P. cubensis and for tolerance to low temperatures,
receiving new breeding materials that contributed to the expan-
sion of germplasm variation. “PlantiCo Zielonki” has a license Academic units
for seed production and sale of the ‘Cezar F1’, which was bred
at InHort. This hybrid is a significant part of “PlantiCo’s” total Both the Warsaw University of Life Sciences and InHort Ski-
sales of cucumber seeds. erniewice are state-sponsored scientific institutions, active main-
ly in fundamental research in their well-equipped laboratories.
“Spójnia Nochowo” They also have greenhouses and experimental fields with an eco-
After the nationalization of the company in the 1950s, logical field in Skierniewice. Both institutions have accumulated
a cucumber for greenhouse production, ‘Nochowski’, was large collections of cucumber cultigens, used also in their applied
maintained (Pędziński 2016). In 1979, a hotbed hybrid ‘Olimp activities such as breeding of new cultivars, to increase the range
F1’ was released and in 1984 another for greenhouse produc- of cucumbers in the domestic market. Both institutions have ex-
tion was released, ‘Lech F1’, which is still active in the regis- tensive cooperation with Polish breeding companies. Cucumber
ter. In the 1980s, thanks to the company president Eng. Mar- is a popular model plant among Polish scholars in fundamental
ian Pędziński, main business areas were formulated; these research. Major players involved in research and breeding are
included breeding of vegetables, seed production, and seed named below, but scientists from Warsaw University, the Polish
marketing. Since then, the company has experienced a marked Academy of Sciences, the University of Łódź, the University of

13
Wrocław, and the Poznań University of Life Sciences have also cucumber haploids and their diploidization (Malepszy 1987).
contributed to various aspects of cucumber biology. Three methods of diploidization of cucumber haploids have been
Cucumber breeding activities in Polish academia are support- developed. The resulting homozygous lines have been applied
ed by various funding sources. So far, the major portion of the in breeding by Polish seed companies. Testing for resistance to
funding was through statutory projects of the Polish Ministry of Pseudomonas syringae pv. lachrymans (angular leaf spot), in co-
Science and Higher Education. In accordance with the gradual re- operation with seed companies, has received Extreme Picture Finder 3.51.4 Crack - Activators Patch atten-
duction of these subsidies in recent years, faculty search to obtain tion (Olczak-Woltman 2008). Currently, cucumber transcriptome
grants from the National Science Center and the National Center analyses in response to infection with this pathogen are under
for Research and Development. Grants awarded by the Polish study. Breeding of cucumber has focused on earliness, yield, and
Ministry of Agriculture and Rural Development (PMARD) have tolerance to angular leaf spot and downy mildew. Much effort
proven particularly important for breeding projects in prepara- has been invested in selecting parental lines for F1 hybrids; this
tion of the starting materials for breeding of vegetable plants and requires knowledge of the genetic basis of parental components,
fundamental research for the biological advancement of plant which is crucial for high seed production efficiency. Over the
production. Plant breeders of InHort have also been carrying out past 25 years, the team has developed a number of cucumber
related applied works under the Multi-Annual Program funded hybrids for field production, the first of which was ‘Dar’ (1993),
by PMARD. Other funds used for the purposes of breeding in- a tribute to the activity of Prof. Kubicki. Subsequently, the team
clude fees from the licenses granted to seed companies. of Prof. Niemirowicz-Szczytt released in 1998-2012: ‘Tessa F1’,
‘Szeryf F1’, ‘Lokata F1’, and ‘Traper F1’. The team also main-
Warsaw University of Life Sciences, Department of Plant tains the cucurbit collection for InHort’s Gene Bank.
Genetics, Breeding, and Biotechnology
From 1965 to 1969, Prof. Kubicki, while simultaneously Institute of Horticulture (former Research Institute
working at the Department of Plant Genetics (Polish Academy of Vegetable Crops) in Skierniewice (InHort; former RIVC)
of Sciences, Skierniewice) and under the leadership of the em- Research on heterosis breeding of cucumber in Skierniewice
inent geneticist Prof. Edmund Malinowski, and part-time at the was initiated and led until 1964 by Barbara Miernik-Pinchinat,
Department of Plant Breeding and Seed Science (RIVC), devel- then continued by Dr. Kłossowska in collaboration with Prof.
oped a method of cucumber F1 hybrid cucumber seed production Kubicki (Malepszy 1987). The resulting breeding materials were
based on gynoecious maternal parents (Kubicki 1969, Niemiro- used to develop the first Polish cucumber, ‘Polan F1’ (Kozik and
wicz-Szczytt 1987, 2016). Together with Dr. Kłossowska, they Doruchowski 2016). Creation of this hybrid resistant against
bred the first Polish hybrid cultivar of cucumber, ‘Polan F1’. C. cucumerinum and CMV was a major breeding achievement,
Together with Dr. Janina Bażant, they developed cultivars for as it yielded two times more than the commonly grown ‘Wisco-
protected cultivation, ‘Iwa’ and ‘Skierniewicki’ (both in 1972); sin SMR-18’ or ‘Monastyrski’ (Kłossowska and Kubicki 1970).
‘Skierniewicki’ and ‘Polan’ are still in the COBORU register. The maternal line was handed over to Polish seed companies.
After the deployment of this method and the maternal lines to “Spójnia Nochowo” used it to breed ‘Śremski F1’. At that time,
Polish breeding stations, Prof. Kubicki supervised the breeding Dr. Kłossowska and Prof. Kubicki initiated testing for resistance
efforts, and became in the 1970s a co-breeder of nine cucumber to angular leaf spot (Kłossowska 1976) and Dr. Janina Bażant
hybrid cultivars. As a member of the faculty of the Department for resistance to powdery mildew (Erysiphe spp.; Bażant 1981).
of Genetics (the name changed several times) at the School of The year 1985 marked the first recorded occurrence of
Horticulture, Warsaw University of Life Sciences, until his un- Pseudoperonospora cubensis in Poland, and since then it has
timely death in 1985, he conducted extensive research on plant been present every year with high severity, often times epidemic.
sexuality, including cucumber. His efforts also focused on het- Prof. Doruchowski’s team reported the mode of inheritance of
erosis and transgressive segregation for earliness, cold toler- resistance to downy milew (Doruchowski and Łąkowska 1992).
ance, pathogen resistance, and nutrient content. Prof. Kubicki Later on, results of parallel studies run in the U.S.A., India, and
also initiated research on interspecific crosses, polyploidy and Poland suggested the possibility that there were different races of
haploidy, and mutations. The results of his research are docu- P. cubensis in these countries. Due to the growing impact of
mented in numerous publications, as listed by Malepszy (1987). downy mildew on cucumber production in our country, Dr. Ur-
Prof. Stefan Malepszy, a geneticist and biotechnologist, szula Kłosińska and I cooperated in screening 1300 cucumber
devoted much time and effort to continue the work of Prof. accessions, leading to the identification of cucumbers with re-
Kubicki. As a great organizer, he continued the construction of sistance levels exceeding those used in previous breeding work
the laboratory, a semi-automatic experimental greenhouse, and (Call et al. 2012). Furthermore, a comprehensive methodology
other infrastructure projects. He also maintained the high level was developed for testing cucumber germplasm under phytotron
of experimental work and directed a number of research topics conditions, using various inoculation methods (Kozik et al.
and projects on cucurbit breeding and genetics. Specifically, 2013). The results correlated with those of field studies under
Prof. Malepszy focused efforts on the effect of genetic back- natural pathogen infection. Resistance in one of the new sourc-
ground on the ability to regenerate plants in vitro, methodolog- es, ‘Ames 2354’, was found to be polygenically inherited. Cur-
ical and biological conditions favoring utilization of genetic rently, the InHort team works toward introgression of genes for
modifications, and production of genetically modified plants of P. cubensis resistance from new sources into horticulturally elite
potential economic value. cucumber germplasm.
Mentoring of the team of vegetable plant breeders has been One result of efforts by Prof. Doruchowski and his collab-
conducted by Prof. Katarzyna Niemirowicz-Szczytt. Under her orators were the first Polish field cucumber F1 hybrids resistant
guidance, the team has continued improving the induction of to Pseudoperonospora cubensis: ‘Aladyn’, ‘Parys’, and ‘Cezar’

14
(1993). They bred a series of F1 cucumbers (‘Atlas’, ‘Pola’, Literature Cited
‘Cyryl’, ‘Bazyl’, ‘Izyd’, ‘Hermes Skierniewicki’, ‘Kronos
Skierniewicki’, ‘Odys’) in the years 1996–2000, having a high Bażant J (1981) Ocena wartości uzytkowej, określenie mechanizmu
level of P. cubensis resistance combined with other useful fea- dziedziczenia odporności na mączniaka prawdziwego Erysiphe ci-
tures. After Prof. Doruchowski’s retirement in 2000, I took over choracearum u nowo wyhodowanej odmiany ogórka do uprawy
the cucumber breeding program together with Dr. Kłosińska, w szklarni i pod osłonami. PhD thesis, Research Institute of Vegetable
and we have succeeded in releasing more field cucumber hybrids Crops, Skierniewice, 1–77.
(‘Rodos’, ‘Reja’, ‘Ozyrys’, ‘Zefir’, ‘Edyp’, ‘Ibis’) with high Call AD, Criswell AD, Wehner TC, Kłosińska U, Kozik EU (2012) Screen-
levels of resistance to P. cubensis. ing for cucumber resistance to downy mildew caused by Pseudoperono-
From 2009 to 2013, my collaborators and I have searched for spora cubensis (Berk. and Curt.) Rostov. Crop Sci 52: 577–592.
Doruchowski RW, Łąkowska-Ryk E (1992) Inheritance of resistance to
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which yielded OPX18950, a marker that distinguished resistant
Cucumis sativus L. In: Doruchowski RW, Kozik EU, Niemirowicz-Szczytt
genotypes from susceptible ones. Subsequently, three quantita- K (eds) Eucarpia Cucurbitaceae ‘92. Skierniewice, pp 132–138.
tive trait loci (QTLs) on cucumber chromosome 5 conferring Karpiński S, Bocian S, Witek Z, Molak D, Marzec L (2016) PlantiCo
downy mildew resistance were mapped and three respective Zielonki. In: Michalik B (ed) Historia hodowli roślin warzywnych
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and molecular mechanisms of resistance. regarding to resistance to angular leaf spot of cucumber. Genet Polon
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self in collaboration with Prof. Todd Wehner of North Carolina Kłossowska E, Kubicki B (1970) Badania nad mieszańcami F1 pomię-
State University from 2003 to 2005, showed that this trait at the dzy różnymi żeńskimi i jednopiennymi odmianami ogórka. Biul Warz
seedling stage was controlled by a single dominant gene, Ch 11: 275–285.
(Kozik and Wehner 2008). In 2014, InHort introduced ‘Ibis F1’, Korohoda J (1984) Rozwój hodowli roślin warzywnych w okresie 40 lat
the first hybrid cucumber with tolerance to low temperatures PRL. Hodowla Roślin 6: 1–26.
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are searching for sources of water-deficit tolerance in cucumber resistance in cucumber seedlings. J Am Soc Hort Sci 133: 225–227.
and investigating the genetic, physiological, and biochemical ba- Kozik EU, Kłosińska U, Call AD, Wehner TC (2013) Heritability and
sis of this trait. genetic variance estimates for resistance to downy mildew in cucum-
ber Ames 2354. Crop Sci 53: 177–182.
Kozik EU, Daemon tools lite 5.0.1 serial number - Free Activators RW (2016) Dzieje hodowli roślin warzywnych
w dawnym Instytucie Warzywnictwa (obecnie Instytucie Ogrodnic-
Conclusions
twa) w Skierniewicach. In: Michalik B (ed) Historia hodowli ro-
ślin warzywnych w Polsce w latach 1958–2015. Kraków: Drukrol,
This overview of the achievements of Polish cucumber pp 61–120 (in press).
breeding is a reminder of the recent and more distant past, Kubicki B (1969) Investigations on sex determination in cucumber
a summary of its current state, and a statement of the future chal- (Cucumis sativus L.). III. Variability of sex expression in monoecious
lenges Extreme Picture Finder 3.51.4 Crack - Activators Patch to be faced by Polish cucumber breeders. Currently, and gynoecious lines. Genet Polon 10: 3–22.
the COBORU Register lists 92 Polish cucumber varieties Malepszy S (1987) Professor Bogusław Kubicki (1933–1985). Genet
(110 total), of which 95% are hybrids; this indicates how much Polon 27: 179–184.
progress has been made in the field of cucumber breeding in Niemirowicz-Szczytt K (1987) Learning and improvement of plants
Poland over less than 100 years, taking into account the histor- - a purpose of the creative research work of Professor Bogusław
ical tragedies of the nation during this time (WWI and WWII). Kubicki. Genet Polon 27: 185–188.
The richness of the germplasm currently in the possession of Niemirowicz-Szczytt K (2016) SGGW Warszawa. In: Michalik B (ed)
Polish cucumber breeders can be expected to provide further Historia hodowli roślin warzywnych w Polsce w latach 1958–2015.
success in cucumber improvement. The efforts of Polish cucum- Kraków: Drukrol, pp 121–163.
ber breeders can be expected to stimulate, within Poland and Olczak-Woltman H, Schollenberger M, Mądry W, Niemirowicz-Szczytt
beyond, innovation in cucumber breeding and seed production, K (2008) Evaluation of cucumber (Cucumis sativus L.) cultivars
and support the implementation of improved cultivars into the grown in Eastern Europe and progress in breeding for resistance to
farming community. angular leaf spot (Pseudomonas syringae pv. lachrymans). Eur J Plant
Pathol 122: 385–393.
Paris HS, Daunay MC, Janick J (2012) Occidental diffusion of cucum-
ber (Cucumis sativus) 500–1300 CE: two routes to Europe. Ann Bot
Acknowledgements
109: 117–126.
Pędziński M (2016) Spójnia Nochowo. In: Michalik B (ed) Historia
The author thanks Prof. Katarzyna Niemirowicz-Szczytt hodowli roślin warzywnych w Polsce w latach 1958–2015. Kraków:
(Warsaw University of Life Sciences), Eng. Marian Pędziński Drukrol, pp 295–331.
(Spójnia Nochowo), Dr. Izabela Żuradzka (POLAN Cracow), Szczechura W, Staniaszek M, Kłosińska U, Kozik EU (2015) Molecular
and Eng. Stanisław Karpiński (PlantiCo Zielonki) for provid- analysis of new sources of resistance to Pseudoperonospora cubensis
ing information on the cucumber breeding programs in their (Berk. et Curt.) Rostovzev in cucumber. Rus J Genet 51: 974–979.
respective institution and companies, respectively. Wojciech Żuradzka I (2016) Polan Kraków. In: Michalik B (ed) Historia hodowli
Szczechura (InHort) is gratefully recognized for retrieval of the roślin warzywnych w Polsce w latach 1958–2015. Kraków: Drukrol,
statistical data. pp 207–274.

15
Emergence of the Sweet Dessert Watermelon, Citrullus lanatus,
in Mediterranean Lands During the Roman Era
Harry S. Paris
Agricultural Research Organization, Newe Ya‘ar Research Center, P. O. Box 1021,
Ramat Yishay 3009500, Israel
e-mail: hsparis@volcani.agri.gov.il

Abstract. Watermelons, Citrullus species, are native to Africa. Archaeological remains, iconography, ancient
literature, and the presence of wild and primitive watermelons in northeastern Africa indicate that the dessert
watermelon, C. lanatus, is native to that region. The dessert watermelon was nurtured and domesticated there, for
water and food, over 4000 years ago. Although the domesticated watermelons of Egypt and Sudan were probably
not bitter, there is no evidence to indicate that they were sweet. Indeed, the extant primitive watermelons known
as gurma in Egypt and gurum in Sudan are spherical and small (≤ 14 cm diameter), with watery but bland, white
or pale green fruit flesh. Hebrew-language literature from the first centuries CE indicates that, by the time of
the Roman Empire, sweet dessert watermelons were esteemed in the Land of Israel, and thus likely were present
in other Mediterranean lands as well. The ripe fruit flesh of the dessert watermelons of that time, which was
probably distinctly colored rather than pale, was eaten raw and had a sweetness which was comparable to that of
figs, grapes, and pomegranates. The seeds were not consumed.
Keywords: Citrullus lanatus, evolution under domestication, watermelon

Introduction However, in spite of the familiarity, dating to 5000 years


Extreme Picture Finder 3.51.4 Crack - Activators Patch ago, of ancient Egyptian civilizations with watermelons,
The dessert watermelon, Citrullus lanatus (Thunb.) Matsum. I could find no evidence showing that the dessert watermelons
& Nakai, is one of the most cooling, refreshing, and appreci- of Egyptian antiquity were sweet. Here I will discuss the evi-
ated food items on hot summer days. Watermelons are among dence for the emergence of sweet dessert watermelons during
the most widely grown vegetable crops in the warmer regions the Roman era.
of the world, with over 3,400,000 hectares planted and over
100,000,000 t harvested annually (Wehner 2008). However, the
sweet dessert watermelons that are so familiar today are derived Materials & Methods
from ancestors that, anthropocentrically, were much inferior.
The xerophytic genus Citrullus (2n = 2x = 22) is native to Sources of evidence
Africa. In accordance with the classification of Chomicki and A multidisciplinary approach that encompasses botany, horti-
Renner (2015), there are seven species in the genus. Three of culture, cookery, philology, and archaeology is necessary to best
them, C. ecirrhosus Cogn., C. rehmii De Winter, and C. naudini- assess crop plant history (Dalby 2003). For cucurbit crops, ar-
anus (Sond.) Hooker f., grow wild in southern Africa and have chaeological remains, iconography, and literature have provided
not been introduced to cultivation. The other four Citrullus spe- much information concerning their development under cultiva-
cies are cultivated to a lesser or greater extent. The colocynth, tion (Paris 2000, Janick et al. 2007).
C. colocynthis (L.) Schrad., is native to northern Africa. The cit-
ron watermelon, C. amarus Schrad., is native to southern Africa. Some diagnostic features of Citrullus
The egusi watermelon, C. mucosospermus (Fursa) Fursa, is na- Citrullus is readily distinguished from other cucurbit genera
tive to western Africa. Recently, I presented a view, based on by the pinnatifid shape of its leaf laminae (Paris et al. 2013). The
archaeological remains, iconography, ancient literature, and flowers are solitary and have five light yellow petals. Most of the
the modern presence of wild and primitive watermelons in Su- flowers are staminate, a pistillate or hermaphroditic flower appear-
dan and Egypt, that the familiar, esteemed dessert watermelon, ing at every seventh or eighth leaf axil (Rosa 1928). Ovaries and
C. lanatus, is native to northeastern Africa (Paris 2015). primordial fruits are lanate, becoming glabrous, smooth, and glossy
as they grow. Usually 30–40 days ensue from anthesis to fruit ma-
turity. External indications of fruit ripening are subtle (Thompson
and Kelly 1957). If stored in a cool, shady place, watermelons can
E.U. Kozik and H.S. Paris (Eds.): Proceedings of Cucurbitaceae 2016,
the XIth EUCARPIA Meeting on Genetics and Breeding of Cucurbitaceae. keep for weeks or even months after harvest without serious dete-
July 24-28, 2016, Warsaw, Poland. rioration of their quality (Keith-Roach 1924, Rushing et al. 2001).

16
Watermelon fruits can weigh as much as 100 kg but most medicine, religion, travel, and cookery. No direct description of
modern, commercially available watermelons range from 3–13 the characteristically smooth, glossy, green-striped rind, or the
kg. Fruit shape can be spherical, globular, oval, or oblong. The taste, texture, or color of the fruit flesh or seeds of watermel-
watermelon rind consists of two layers. The surface of the thin, ons has been found in literature of antiquity (Janick et al. 2007,
glossy exocarp is typically boldly striped in two shades of green; Paris 2015).
the stripes are jaggedly edged and range in breadth from very Caution must be exercised in interpretation of the ancient
narrow to very broad. The thick, white mesocarp is wet and hard. literature. Adjectives tend to be used differently across languag-
Underneath the rind is the watery fruit flesh or endocarp, which es, contexts, geographic areas, and periods of time (Paris et al.
is the portion of the fruit that is usually eaten. Early in devel- 2012). For example, “sweet” can be synonymous with sugary or
opment, the fruit flesh is hard, white or otherwise pale-colored, not bitter, not sour, not salty, or not spicy. Also, what might have
and insipid. In citron watermelons, the fruit flesh remains hard, been considered sweet in ancient times might not be considered
nearly colorless and tasteless to fruit maturity (Xu et al. 2012). sweet today, given recent development of cultigens with greater
In sweet dessert watermelons, the flesh of the maturing fruit be- sweetness. The adjective “red” has been variously used for or-
comes soft and accumulates carotenoid pigments and sucrose ange, purple, and brown, and “yellow” has been used for orange.
(Elmstrom and Davis 1981, Brown and Summers 1985, Soteriou Nouns have also been used indiscriminantly. The word “melon”
et al. 2014). Color begins to accumulate between 2 and 3 weeks has been used in American English for both Citrullus and Cucum-
after anthesis, first around the developing seeds and thereafter is melo. Total uninstaller 3.3.0.161 license key examples occur for the medieval Latin pepo and
gradually spreading throughout the endocarp (Perkins-Veazie melones, and Arabic battikh. The intention of such words has to
et al. 2012). Depending on the genotype, the flesh color of ripe be interpreted in the context of time period, language, and geo-
watermelon fruits can range from red, pink, orange, yellow, or a graphic area.
mixture of these colours, to green or white (Gusmini and Wehner
2006). The range in texture of the ripe fruit flesh can be described
as crisp, tender, or liquefied, and coarsely or finely grained. Each Watermelons in Literature of Antiquity
fruit can contain several hundred seeds that, to the casual observer,
are scattered throughout the flesh and, to the consumer, are of Biblical Hebrew
much annoyance. The seeds of dessert watermelons are hard, The Children of Israel, during their sojourn in the Sinai Des-
flat, and oval and, depending on the genotype, range in length ert after the exodus, longed for five vegetables they knew from
from 10 to 16 mm and are black, brown, tan, white, yellow, or the Land of Egypt; these were, respectively, the qishu’im (veg-
red, and can be patterned with a second color. etable melons), avattihim (watermelons), hazir (leeks), bezalim
Dessert watermelons, C. lanatus, are sometimes confused (onions), and shumim (garlics) (Numbers 11:5). As watermel-
with melons, Cucumis melo L., as both are often large and sweet. ons were mentioned together with other vegetables, they were
The most salient features distinguishing the two are the shape of considered to be just like the others, to be eaten raw, cooked,
the leaf laminae, distribution of staminate and pistillate flowers or pickled. They were not like modern sweet dessert watermel-
on the plant, range of fruit shape, fruit surface features, wetness ons, otherwise they would have been craved to be eaten first and
of the fruit, thickness of the fruit rind, fruit flesh color, and shape, immediately in a desert environment.
color, and distribution of seeds within the fruit (Paris et al. 2012). Centuries later, in the Land of Israel, the word used for
In the field, watermelons ripen evenly over the course of the a field of cucurbits was miqsha (Isaiah 1:8). The word miqsha is
harvest season but melons ripen in distinct waves (Rosa 1924, derived from the word qishu’im. Indeed, in those times, water-
McGlasson and Pratt 1963, Pratt et al. 1977). Watermelons have melons must have been of secondary importance to the vegetable
no well-marked indicators of fruit ripening whereas melons typ- melons, and there is no evidence suggesting that the watermelons
ically become aromatic and yellow, and abscise from the plant were sweet.
upon ripening (Isenberg et al. 1987, Nonnecke 1989). Water-
melons have a much longer shelf-life than most melons but are Greek (400 BCE – 355 CE)
subject to breakage if not handled properly (Whitaker and Davis apowermirror vip crack mac - Activators Patch The word pepon of classical Greek, literally a sun-ripened
1962, Robinson and Decker-Walters 1997). In watermelons, the fruit, usually referred specifically to watermelon (Liddell and
seeds are distributed within the fruit flesh but in melons the fruit Scott 1948, Andrews 1958, Stol 1987, Grant 2000). The Greek
flesh is free of seeds. doctor Hippocrates, in approximately 400 BCE, wrote that
the pepones were easily digestible (Jones 1967). Likewise,
Possible pitfalls in interpretation of literature Dioscorides, in 70 CE, wrote that the flesh of the pepon is easily
Although iconography has been the most important source digestible and diuretic, and prescribed the rind of the pepon to
of evidence in understanding cucurbit crop history (Eisendrath be applied on top of the head, for children suffering from heat
1961), fruit sweetness is not a trait that is readily amenable to stroke (Osbaldeston and Wood 2000, Beck 2005). A century
illustration or detection in archaeological remains. Literature later, Galen wrote that the pepon was cold and wet, and a diuret-
is a potentially rich source of information concerning possi- ic, and the melopepon less so (Grant 2000, Powell and Wilkins
ble sweetness of cucurbit fruits (Paris et al. 2012). Generally, 2003). Clearly, to all of these doctors, the pepo they were
though, food items were considered by ancient writers to be fa- describing was the watermelon, C. lanatus. The melopepon, in-
miliar to everyone, and thus in no need of description, and were troduced later and of lesser effect, would appear to be the melon,
usually discussed only in reference to their supposed dietary or C. melo.
medicinal effects (Dalby 2003). Off-hand, indirect descriptions Athenaeus, in The Learned Banqueters (late second century),
of foods can sometimes be gathered in other contexts, especially quoted thousands of lines of verse written by approximately

17
1000 Greek authors from various times and localities (Olson an intermingling. This is called in the Greek language molefe-
2006). One of them, Phaenias, is quoted as stating that the pepon fon.” Thus the myth of the apple-pepo was carried across three
is edible, except for the seeds, when the flesh becomes soft. An- languages, Greek, Latin, and Hebrew. More importantly, it al-
other, Diocles, wrote that the best kolokyntas were round, very lows definite identification of the Hebrew melofefonot as melons,
large, glukeian (sweet), and easy to digest. The word kolokyntas C. melo. Another passage from the Jerusalem Talmud (Kil’ayim
was usually applied to bottle gourds, Lagenaria siceraria (Mol.) 1:8, 4a) has this comment: “Prohibited is the insertion of cut-
Standl., but for some ancient Greek speakers the word was ap- tings from grapevines into an avattiah lest it throw its waters into
plied to watermelons. them.” The avattiah, like the Greek pepon and Latin pepo, was
a large, watery fruit, the watermelon, C. lanatus.
Latin (77 – 516 CE) The four edible-fruited cucurbits are considered togeth-
Pliny, in his Historia Naturalis (77 CE), described the pepo er in the first chapter of the tractate on tithing, Ma‘asrot. The
as a most refreshing and cooling food (Rackham 1950, Jones qishu’im and delu‘im were to undergo piqqus, rubbing off of
1951). This description would fit the watermelon, C. lanatus. their hairs, prior to eating. Hence, these two cucurbit fruits, vege-
The melopepo (apple-pepo) was a new introduction resembling table melons and bottle gourds, were eaten young and immature,
a quince that, upon ripening, turned yellow, became aromatic, when they were hairy. In contrast, the avattihim were to be tithed
and, significantly, spontaneously detached from the plant. Ob- after they underwent shilluq, smoothing or polishing for removal
viously, the melopepo was a cultigen of melon, C. melo. Quin- of the dust that accumulated on them. Therefore the avattihim,
tus Gargilius Martialis, around 260 CE, wrote that the pepone watermelons, were eaten when ripe. Corroborating evidence for
were good to eat after removal of the rind and pits (Maire 2007). the eating of the watermelons when they were ripe is also found
De Observantia Ciborum (510 CE) has a list 101 foods, includ- in the Mishna tractate on peduncles (‘Oqazin 2:3), in which it is
ing three cucurbits (Mazzini 1984). The pepone are not listed mentioned that a partially withered pomegranate or watermel-
with the other two cucurbits. Instead, they are among pome- on is not eaten. The tractate on tithing in the Tosefta indicated
granates, grapes, and figs, which are sweet, juicy fruits that that the melafefonot, melons, were to be tithed after they were
are usually eaten raw when ripe. Anthimus, in De Observantia taken out of the field and gathered (Ma‘asrot 1:6). Specifically,
Ciborum Epistula (516 CE), indicates that the melones were to the grower was obliged to tithe each wave (harvest) of ripened
be eaten well-ripened, fresh with the seeds still mixed in the melons.
flesh (Grant 2007). malwarebytes 4.2.0 license key 2020 - Free Activators Melon seeds were to be tithed but watermelon seeds were not,
because the latter could not be used as food (Jerusalem Talmud,
Hebrew (150 – 400 CE) Ma‘asrot 1:2, 2b). This might at first seem strange as, today, wa-
Three large codices of Jewish Law were compiled in Israel termelon seeds are consumed in many areas. However, watermel-
during the first centuries CE. They are easily searched at Mekhon on seeds of the time were quite small, only 11 mm long (Cox and
Mamre (http://www.mechon-mamre.org). The first of these is the van der Veen 2008, Kislev and Simhoni 2009), and given their
Mishna, which was compiled during the latter half of the second thick, hard seed coats were not particularly suitable for use as
century. The second, the Tosefta, was compiled approximately food.
a century later and differs in some of its material from the Mish- The tractate on contributions, Terumot, contains a discus-
na. The third, the Jerusalem Talmud, probably compiled during sion on the amount of time, post-harvest, that various fruits and
the latter half of the fourth century, includes a text very similar to vegetables are acceptable as contributions. As would be expect-
that of the Mishna but with much exposition. All three codices are ed from highly perishable, young cucurbit fruits, the vegetable
arranged in sedarim (orders) each of which has varying numbers melons and bottle gourds were suitable for contribution for only
of massakhtot (tractates). The Mishna has 63 tractates sorted into one day after harvest (Tosefta, Terumot 4:5). The melons were
six orders. Extreme Picture Finder 3.51.4 Crack - Activators Patch acceptable for three days. The watermelons are not mentioned
Some of the tractates refer to edible-fruited cucurbits, most in this context, apparently because they had a long shelf life.
notably the ones on Kil’ayim (Intermingling), Ma‘asrot (Tith- Also unlike the other three cucurbits, watermelons could not be
ing), Terumot (Contributions), and ‘Oqazin (Peduncles). Four gathered for sale in a pile. Instead they had to be laid out one-by-
edible cucurbits are often mentioned, these four in Hebrew being one (Jerusalem Talmud, Ma‘asrot 1:4, 4a), indicating that they
the qishu’im (vegetable melons, Cucumis melo), delu‘im (bottle were fragile, and thus very different from the modern cultivars of
gourds, L. siceraria), avattihim, and melafefonot. A fifth edible Extreme Picture Finder 3.51.4 Crack - Activators Patch that have been bred for adaptation to long-distance
cucurbit, the qarmulin, is mentioned only in the latter two works, shipping.
apparently being a new introduction with an appearance and Finally and most significantly, watermelons differed in one
use similar to the delu‘im. The qarmulin have been identified as more way from the other edible cucurbits. In the tractate on tith-
sponge gourds, Luffa aegyptiaca Mill. (Janick et al. 2007, Avital ing, watermelons are discussed together with figs, table grapes,
and Paris 2014). and pomegranates. Fruits that were picked in the garden or field
The four edible-fruited cucurbits are considered together in and eaten there were exempt from tithing (Mishna, Ma‘asrot 2:6;
the tractate Kil’ayim because of the concern given to their vines Jerusalem Talmud, Ma‘asrot 2:4, 11a). Figs were simply cho-
intermingling with one another. There are some elements of sen and eaten, grapes were picked one-by-one from a cluster and
practicality in this prohibition. On the other hand, in the Jerusa- eaten, pomegranates were plucked and eaten, and watermelons
lem Talmud version (Kil’ayim 1:2, 2a), there is also a recantation were sliced and eaten. Evidently, the watermelons of the latter
of Mediterranean agricultural folklore: “A person takes a seed half of the second century in Israel, like figs, table grapes, and
from the flesh of an avattiah and a seed from the flesh of an apple pomegranates, were common fruits that were eaten raw without
and puts them together in the same hole and they unite to become any culinary preparation and, like them, were juicy and sweet.

18
Discussion in Khorasan, Central Asia, during early medieval times, reaching
Mediterranean lands in the eleventh century (Paris et al. 2012).
Domesticated plants are derived from small samples of wild The Hebrew evidence for the existence of sweet dessert wa-
source populations, and thus are themselves founder populations termelons in Mediterranean lands by Roman times is corrob-
that contain only a small fraction of the genetic diversity of their orated by a contemporary Greek writing and a Latin writing
wild ancestors (Ladizinsky 1998). Cultigens have various traits from just after the fall of the Roman Empire. In the late sec-
that were selected early and continually in the domestication pro- ond century, Athenaeus quoted Diocles as stating that the
cess, such as lack of bitterness, increased size of the harvested best kolokyntas are glukeian, sweet (Olson 2006). The early
parts, increased yield, and novel coloration (Heslop-Harrison and sixth-century Latin De Observantia Ciborum lists the pepone
Schwarzacher 2012). The sweet dessert watermelon, C. lanatus, among pomegranates, grapes, and figs, confirming the presence
which has relatively little genetic diversity (Levi et al. 2001, of sweet watermelons in what is now Italy (Mazzini, 1984).
Reddy et al. 2015), follows this general pattern of crop-plant do- Thus, sweet watermelons had diffused to Europe centuries be-
mestication. The fruit flesh of wild and primitive Citrullus is bit- fore their earliest recorded presence in Moorish Spain, in the
ter or insipid, hard, and pale-coloured (Wehner 2008). Non-bit- Cordoban Calendar of 961 CE (Pellat, 1961).
terness of the fruits was probably the first and most important
trait to be selected in the process of watermelon domestication.
As non-bitterness is conferred by a single recessive gene (Weh- Conclusion
ner 2007), this trait should have been relatively easy to maintain
if isolation from neighbouring wild populations was feasible. Sweet dessert watermelons were selected from non-sweet an-
The modes of inheritance of hard versus soft and of insipid cestors. By the time of the Roman Empire, sweet dessert water-
versus sweet flesh in Citrullus have not been illuminated. melons were a familiar and esteemed produce item in Israel, and
Although it is agreed that the genus Citrullus is of African ori- probably in other Mediterranean lands as well.
gin, there has been major disagreement concerning where in Africa
the dessert watermelon originated. Much of the controversy stems
from phenotypic variations that overlap among C. colocynthis, Acknowledgements
C. amarus, C. mucosospermus, and C. lanatus, and the weak
crossability barriers among these four species. Regardless of Financial support of the Lillian Goldman Charitable Trust
species, wild and primitive Citrullus fruits typically have hard, (New York) is gratefully acknowledged.
bitter or bland, weakly coloured flesh and, as indicated by Weh-
ner (2008), this situation has repeatedly been a source of incor-
rect taxonomic identifications. Archaeological remains, iconog- Literature Cited
raphy, and literature, as well as the modern presence of wild and
primitive watermelons in Sudan and Egypt, support the idea Andrews AC (1958) Melons and watermelons in the classical era. Osiris
that the dessert watermelon, C. lanatus, is native to northeastern 12: 368–375.
Africa (Paris 2015). Avital A, Paris HS (2014) Cucurbits depicted in Byzantine mosaics
The present investigation, which has focused mainly on an- from Israel, 350–600 CE. Ann Bot 114: 203–222.
cient literature, has attempted to define a narrower time frame Beck LY (2005) Pedanius Dioscorides of Anazarbus: De Materia Medi-
and geographical range within which the dessert watermelons ca. Hildesheim: Olms–Weidmann.
having non-bitter, tender, highly coloured, sweet flesh were de- Brown AC, Summers WL (1985) Carbohydrate accumulation and color
veloped. The latest possible date for this time frame is provid- development in watermelon. J Am Soc Hort Sci 110: 683–686.
Chomicki G, Renner SS (2015) Watermelon origin solved with mo-
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lecular phylogenetics including Linnean material: another example
and white-flesh citron watermelons in illustrated manuscripts
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20
Status of Cucurbit Breeding at AVRDC
– The World Vegetable Center
Narinder P.S. Dhillon1, Supannika Sanguansil1, Supornpun Srimat1,
Hsin-chun Cheng2, Chung-cheng Lin3, Ramasamy Srinivasan4, Lawrence Kenyon5,
Roland Schafleitner6, Ray-yu Yang7, and Peter Hanson2
1
AVRDC – The World Vegetable Center, East and Southeast Asia, Kasetsart University, Kamphaeng Saen,
Nakhon Pathom 73140, Thailand
2
Breeding, AVRDC – The World Vegetable Center, P.O. Box 42, Shanhua, Tainan 741, Taiwan
3
Breeding, AVRDC – The World Vegetable Center, P.O. Box 42, Shanhua, Tainan 741, Taiwan;
Present address: Clover Seed Company Ltd., Hong Kong Village Lots 91-104, Snowsonhill,
Hong Kong, P.R. China
4
Entomology, AVRDC – The World Vegetable Center, P.O. Box 42, Shanhua, Tainan 741, Taiwan
5
Virology, AVRDC – The World Vegetable Center, P.O. Box 42, Shanhua, Tainan 741, Taiwan
6
Biotechnology, AVRDC – The World Vegetable Center, P.O. Box 42, Shanhua, Tainan 741, Taiwan
7
Nutrition, AVRDC – The World Vegetable Center, P.O. Box 42, Shanhua, Tainan 741, Taiwan
e-mail: narinder.dhillon@worldveg.org

Abstract. AVRDC – The World Vegetable Center’s global cucurbit breeding program started in 2006 at Center
headquarters in Taiwan. The program shifted to AVRDC East and Southeast Asia, Thailand, in May 2010.
Breeding efforts are focused on two cucurbit species, bitter gourd (Momordica charantia L.) and tropical pumpkin
(Cucurbita moschata Duchesne), with comparatively less emphasis on cucumber (Cucumis sativus L.). The Center’s
bitter gourd breeding efforts have resulted in the development of high yielding open-pollinated lines for different
market segments with superior fruit quality, early maturity, ability to set fruit under high temperatures, high
concentration of anti-diabetic compounds, and disease and insect resistance. Eight bitter gourd open-pollinated lines
of different market segments with yield and fruit quality comparable to commercial hybrids have been released.
Bitter gourd lines resistant to powdery mildew (Podosphaera xanthii) have been developed and the resistance has
been confirmed in multi-country trials in Asia. Bitter gourd lines resistant to leaf spot (Cercospora spp., Alternaria
spp.) and anthracnose (Colletotrichum spp.) have been identified. Melon fly (Bactrocera cucurbitae) resistance in
Momordica balsamina L., a close relative of the bitter gourd, has been identified. Populations for further selection
through inbreeding for improved yield, fruit quality, and disease resistance have been created at AVRDC for
various market segments by crossing genetically unrelated bitter gourd varieties and genebank accessions. Four
cycles of recurrent selection have been completed. Pumpkin breeding at AVRDC has led to the development of
open-pollinated lines rich in beta-carotene with good eating quality and resistance to Cucumber mosaic virus
(CMV), Zucchini yellow mosaic virus (ZYMV), Papaya ringspot virus – watermelon strain (PRSV-W), and Squash
leaf curl virus (SqLCV). AVRDC’s cucumber breeding program has developed non-bitter, predominantly female,
high yielding tropical lines suitable for the Southeast Asian market.
Keywords: Momordica charantia, Cucurbita moschata, Cucumis sativus, breeding, disease/insect-pest resistance,
phytonutrient

E.U. Kozik and H.S. Paris (Eds.): Proceedings of Cucurbitaceae 2016,


the XIth EUCARPIA Meeting keyword researcher pro review - Free Activators Genetics and Breeding of Cucurbitaceae.
July 24-28, 2016, Warsaw, Poland.

21
Introduction dery mildew caused by Podosphaera xanthii is a serious fun-
gal disease of bitter gourd, and yield losses of up to 50% have
AVRDC’s cucurbit breeding program was started in 2006 at been observed in China and India (Fu Jiqin, Enza Zaden,
the Center’s headquarters in Taiwan and shifted to AVRDC East Vimal Chawda, VNR Seeds Pvt. Ltd., personal communica-
and Southeast Asia, Thailand in May 2010. Considerations to Prima Cartoonizer 1.6.5 Crack + Keygen Key Free Download 2020. The disease can be controlled with fungicides, but ge-
select the target cucurbit crops included economic importance, netic resistance provides a more economically sound and en-
nutrient density, access to genetic resources, and AVRDC’s com- vironmentally safe approach. After screening 150 accessions
parative advantage versus the private sector. The cucurbit breed- of a global collection of bitter gourd against the local isolate of
ing program initially emphasized improvement of cucumber, P. xanthii at Kamphaeng Saen (Thailand), the cucurbit breeding
Cucumis sativus L., because of the crop’s economic importance, program developed five inbred lines resistant to this pathogen.
and of tropical pumpkin, Cucurbita moschata Duchesne, which A single resistant plant was identified from five segregating pop-
is nutritionally rich but has received relatively less attention from ulations derived from genebank accessions; multiple cycles of
the private seed sector. Subsequently, the Center downgraded inbreeding and selection led to development of five resistant in-
cucumber breeding and took up improvement of bitter gourd, bred lines (THMC 113, THMC 143, THMC 153, THMC 167,
Momordica charantia L., a crop that is economically and nutri- and THMC 170). These lines were screened against local isolates
tionally important. of P. xanthii at 11 locations in six countries (China, Thailand,
India, Taiwan, Vietnam, Philippines). THMC 153 was resis-
tant across all locations, and THMC 143 and THMC 167, were
Bitter Gourd Breeding at AVRDC resistant at all locations except one in China. The other bitter
– The World Vegetable Center gourd lines exhibited variable reactions to the pathogen in dif-
ferent regions, which confirms the presence of genetic variants
Bitter gourd is an important cucurbitaceous cash crop culti- (pathotypes and physiological races) of P. xanthi with different
vated mainly by smallholder farmers in Asia, where it is grown abilities to infect and sporulate on bitter gourd. THMC 143,
on approximately 340,000 ha annually (McCreight et al. 2013). THMC 153, and THMC 167 have the potential to be sources
Bitter gourd fruits are high in nutrients such as beta-carotene, of broad spectrum disease resistance for breeding purposes. The
vitamin C, folic acid, magnesium, phosphorus, and potassi- resistance of these bitter gourd lines to the Kamphaeng Saen
um (Yuwai et al. 1991, Dhillon et al. 2016). It is often used (Thailand) isolate of P. xanthii is controlled by recessive poly-
in folk medicine to manage Type 2 diabetes, a rapidly spread- genes (N.P.S. Dhillon, unpublished data).
ing non-communicable disease that affects 347 million people The melon fly, Bactrocera cucurbitae (Coquillett) (Dip-
worldwide, with 80% of them living in low-income and mid- tera: Tephritidae) is an important insect pest of bitter gourd and
dle-income countries (WHO 2015). The current bitter gourd other cucurbits. Fruit infestation by this pest has been report-
seed market in Asia is valued at about 16 million Euros (Naren- ed to vary between 41 to 89% in Asia (Rabindranath and Pil-
dra Kumar Singh, HM Clause, personal communication). India lai 1986) and up to 95% bitter gourd fruit infestation has been
is the largest producer of bitter gourd and approximately 350 t reported in Papua New Guinea (Hollingsworth et al. 1997).
of seed (hybrid seed = 200 t and open-pollinated seed = 150 t, Chemical control of this pest is expensive and represents a risk
M. Anil, Rasi Seeds Pvt. Ltd., personal communication) is pro- to the environment. In addition, this insect can rapidly develop
duced annually for the Indian bitter gourd seed market. By 2020, resistance to chemical pesticides. Host-plant resistance is safe,
the market is expected to grow to 600 t (hybrid seed = 360 t, economical, and an effective alternative to chemical pesticides.
open-pollinated seed = 240 t, Vimal Chawda, VNR Seeds Pvt. In a review, Dhillon et al. (2005) listed several cultivated and
Ltd., personal communication). wild bitter gourd accessions as resistant or highly resistant to
Based on a literature survey, Léon (1994) suggested that B. cucurbitae. ‘Faisalabad Long’ and ‘Col-II’, originally derived
hybrids of out-crossing crops had greater yield stability. Bitter from bitter gourd landraces, were resistant to local strains of
gourd displays high heterosis for yield ranging from 27 to 86% B. cucurbitae in Pakistan (Gogi et al. 2010), but these two culti-
above the high parent (Behera 2004). This has encouraged the vars were highly susceptible to the local strain of B. cucurbitae in
development and adoption of hybrid cultivars in Asia, which Taiwan (Table 2). However, THMC 281, an accession of Momor-
have the potential to increase profitability of seed companies dica balsamina L., which is closely related to M. charantia, was
as well as growers. However, approximately 57% of the bit- resistant to the Taiwan strain of melon fly in two field screenings
ter gourd area is planted to open-pollinated cultivars in Asia during 2012 and 2013. Usually, melon fly-resistant bitter gourd
(Ajay Dyal, Rasi Seeds Pvt. Ltd., personal communication). fruits are thick with a tough rind, like the fruits of THMC 281.
Average yield of popular open-pollinated cultivars rang- Hence, it might be worthwhile investigating the basis and mech-
es between 12 to 15 t·ha-1, whereas commercial hybrids yield anisms of melon-fly resistance in this accession, and whether it
20 to 25 t·ha-1 (S.R. Verma, VNR Seeds Pvt. Ltd., Chaiwat could be incorporated into resistance breeding programs.
Manoworn, Chia Tai Seeds, personal communication). AVRDC Feeding studies on animals and humans confirmed that bitter
has developed bitter gourd inbred lines comparable in yield and gourd whole fruit, juice, or extract can play a role in managing
fruit quality to popular commercial hybrids (Table 1). Two of Type 2 diabetes (Krawinkel and Keding 2006). The antidiabetic
these lines (AVBG1310 and AVBG1313) produced stable yields effect of bitter gourd is due to the complex action of multiple
in trials at Kamphaeng Saen (Thailand) in 2013 and 2014. compounds in the fruit. The AVRDC-led research project “A bet-
This result was despite the significantly higher temperatures ter bitter gourd: Exploring bitter gourd (Momordica charantia)
(32 to 38 °C) during the flowering and fruiting stages in 2014, to increase incomes, manage type 2 diabetes, and provide health
compared to the same period in 2013 (28 to 35 °C). Pow- in developing countries” funded by the Federal Ministry for

22
Economic Cooperation and Development (BMZ), Germany, led table Center, Taiwan has a fairly large collection of bitter gourd
to the identification of bitter gourd lines with high and stable germplasm from more than 15 countries. The genetic diversity
concentrations of antidiabetic compounds, and an assessment of 114 bitter gourd accessions of the AVRDC collection and of
of the contribution of these antidiabetic effects to human health commercial cultivars was assessed based on the genotypes at 50
where bitter gourd is consumed as a part of a balanced diet. simple sequence repeat (SSR) loci. Commercial cultivars of sim-
A total of 31 single/groups of compounds from LC/ToF-MS ilar market segments were closely related, even though they were
(Liquid chromatography/Time-of-flight mass spectrometery) released by different seed companies, corroborating the assump-
profiles were identified with a molecular weight (M+H±0.05 tion that the genetic base of bitter gourd varieties in Asia is very
Da) corresponding to triterpenoids (Yang et al. 2015), and 16 narrow. South Asian accessions originating from India, Bangla-
antidiabetic related compounds including seven phosphatidyl desh, and Pakistan were more closely related to each other than
cholines, two fatty acid derivatives, five triterpenoids, vicine videosolo video converter ultimate portable - Free Activators to accessions of any other geographic group. Likewise, acces-
and tryptophan were identified in the LCMS (Liquid chroma- sions from Cambodia, Vietnam, and The Philippines clustered
tography/mass spectrometery) profiles of methanol extracts of into one group, and were distinct from the Taiwan accessions.
M. charantia. Preliminary results of feeding studies with humans There is a pressing need to methodically evaluate bitter gourd
conducted by AVRDC in India revealed that the mean initial fast- accessions held by various national and international genebanks
ing blood glucose levels of pre-diabetics had reduced significant- for resistance against economically important pests and patho-
ly (P < 0.01) after consuming bitter gourd for an extended time. wirecast pro 13 free download - Free Activators gens, assess their tolerance to abiotic stresses, and evaluate their
For the last two decades, bitter gourd breeders have pursued phytonutrient contents and profiles for micronutrients and anti-
genetic improvement of the crop through a process of inbreeding diabetic compounds. For instance, Cucurbit aphid-borne yellow
and pedigree selection to develop lines, followed by elite × elite virus also known as Namamarako in the Philippines and Mara
crosses and identification of commercilizable hybrids. This ap- Ba in Thailand, is an aphid-transmitted polerovirus that has be-
proach was adopted to increase the chances of developing lines come a serious production limitation in bitter gourd (Relevante
possessing multiple important traits preferred by growers, such windows 8.1 crack - Free Activators al. 2012, unpublished). Bitter gourd accession VI049946 from
as earliness and high marketable yield, and consumer-preferred the AVRDC genebank has segregated for resistance to this virus
traits, such as fruit color, shape, size, and skin pattern. Perfor- (Fatkhu Rokhman, East-West Seed Indonesia, personal commu-
mance-driven selection to achieve the preferences of value chain nication) and the company is using selections from this accession
actors (growers, distributors, retailers, and consumers) led to re- to eventually develop resistant cultivars.
peated use of elite inbred lines for hybrid development, which
narrowed the genetic base of the crop and has led to low genetic
diversity among commercial cultivars (Duvick 2005). Lack of Pumpkin Breeding at AVRDC
genetic diversity can enhance vulnerability of the crop to disease – The World Vegetable Center
and insect outbreaks (Keneni et al. 2012). Breeders’ greater use
of the bitter gourd diversity stored in genebanks could eventu- Tropical pumpkin, Cucurbita moschata, is an underutilized
ally broaden the genetic base of elite cultivars and reduce the cucurbitaceous crop with economic potential and nutritional im-
chances of catastrophic outcomes. AVRDC – The World Vege- portance. Pumpkin fruits are rich in alpha and beta-carotenes and

Table 1. Mean fruit number/plant, fruit weight and yield, and fruit traits of AVRDC bitter gourd lines.
Fruit Fruit Yield Fruit Fruit Fruit
No.* Genotype
number/plant weight (g) (t/ha) bitterness color skin pattern
1 AVBG1304 41 173 35.9 Low Dark green Spiny
2 AVBG1310 23 266 33.6 Low Green Spiny
3 Palee (Check) 30 194 36.3 Low Medium green Spiny
4 BARI Karela 1 (Check) 22 178 23.5 Medium Medium green Spiny
5 AVBG1311 18 326 39.4 Low Light green Ridged
6 AVBG1313 19 374 41.3 Low Light green Ridged
7 AVBG1314 21 350 41.3 Low Light green Ridged
8 Benteng (Check) 19 407 40.6 Low Light green Ridged
9 AVBG1323 47 138 32.8 Medium Green Spiny
10 AVBG1324 59 115 38.6 Medium Medium green Spiny
11 AVBG1325 52 129 36.9 Low Green Spiny
12 AVBG1301 45 125 34.9 Medium Medium green Spiny
13 Noor (Check) 48 134 36.3 Low Medium green Spiny
LSD (P = 0.05) 9 70 7.0
*Market segment: 1-4: medium fruit length segment (South Asian type), 5 to 8: long fruit segment (Southeast Asian/Chinese
type), 9 to 13: small fruit length segment (South Asian type).

23
minerals (Arima and Rodriguez-Amaya 1990). The world pro- Future of AVRDC’s Cucurbit Breeding Program
duction of squash, pumpkins, and gourds in 2012 was estimated
at 24.6 million t from 1.78 million ha (FAO 2012). Productivity We are in the process of developing markers to map powdery
of these cucurbits is high in North America (23.7 t·ha-1) com- mildew resistance genes. Currently more than 5,000 polymor-
pared to Asia (13.8 t·ha-1) and Africa (7 t·ha-1). Open-pollinated phic markers have been obtained for a segregating bitter gourd
cultivars still dominate commercial acreage of pumpkin in the population and mapping of the resistance genes is ongoing. Pop-
tropics. Genetic uniformity and disease susceptibility are two ulations of pumpkin are being developed to study the inheritance
major constraints leading to low pumpkin yields in the tropics of resistance to ZYMV, PRSV-W, and SqLCV and molecular
(Loy 2011). Viral diseases caused by Cucumber mosaic virus markers associated with these resistance gene(s). We are test-
(CMV), potyviruses such as Papaya ringspot virus-watermelon ing the performance of AVRDC-bred improved lines of bitter
strain (PRSV-W), Zucchini yellow mosaic virus (ZYMV), and gourd and pumpkin in various countries in Southeast Asia and
begomoviruses such as Squash leaf curl virus (SqLCV) and To- South Asia. We are exploring the potential of extending testing
mato leaf curl New Delhi virus (ToLCNDV), are diseases of ma- of AVRDC bitter gourd and pumpkin lines in parts of Africa.
jor economic importance in the tropics and can cause losses of We are also screening AVRDC genebank accessions to find suit-
up to 100%. Use of host plant resistance is the cheapest and most able prospects for the development of bitter gourd lines resistant
environmentally sound means to control viruses. to Cercospora spp., downy mildew, and begomoviruses. AVRDC
The aphid-transmitted ZYMV is a major pathogen of pump- cucurbit lines have been shared with breeders from private seed
kin. C. moschata landrace Nigerian Local carries a single dom- companies, who have developed hybrids using this breeding
inant resistance gene to the local isolate of ZYMV (ZYMV-CT) material; these hybrids are being tested at multiple locations by
in the U.S.A. (Munger and Provvidenti 1987, Brown et al. product development teams from seed companies (M. Anil, Rasi
2003). Two genes (Zym-0 and Zym-4) for resistance to a mix- Seeds Pvt. Ltd., India; Parag Agarwal, VNR Seeds Pvt. Ltd., and
ture of five Austrian isolates of ZYMV have been identified in Murali Kumar, Sakata Seeds, India, personal communication).
Nigerian Local (Pachner et al. 2011). A recessive gene carried
by ‘Waltham Butternut’, zym-5, is complimentary with the dom-
inant Zym-4 of Nigerian Local and the resistance conferred by Literature Cited
Zym-4 is only expressed in zym-5/zym-5 individuals. We found
that resistance in Nigerian Local to the most virulent strain, Arima HK, Rodriguez-Amaya DB (1990) Carotenoid composition and
ZYMV-TN3, which is prevalent in Taiwan, is controlled by vitamin A value of squash and pumpkin from northeastern Brazil.
two complementary dominant genes. We transferred ZYMV Arch Latinoam Nutr 40: 284–292.
resistance from Nigerian Local into the susceptible local variety Behera TK (2004) Heterosis in bitter gourd. J New Seeds 6: 217–221.
PY through backcross breeding and obtained two pumpkin lines Brown RN, Bolanos-Herrera A, Myers JR, Jahn MM (2003) Inheritance
completely resistant to ZYMV; these lines are comparable in of resistance to four cucurbit viruses in Cucurbita moschata. Euphyt-
yield and fruit quality to a popular local variety and to a com- ica 129: 253–258.
mercial hybrid susceptible to the virus. Pumpkin crops grown Dhillon MK, Singh R, Naresh JS, Sharma HC (2005) The melon fruit
Extreme Picture Finder 3.51.4 Crack - Activators Patch fly, Bactrocera cucurbitae: A review of its biology and management.
in farmers’ fields in the tropics can be infected with multiple
J Insect Sci 5: 40.
virus species and cultivars carrying resistance to only a single Dhillon NPS, Lin CC, Sun Z, et al. (2016) Varietal and harvesting stage
virus will not perform well. Potyviruses and begomoviruses are variation in the content of carotenoids, ascorbic acid and tocopherols
present at high incidences in the pumpkin fields at the Research in the fruit of bitter gourd (Momordica charantia L.). Plant Genet Re-
and Training Station of AVRDC – The World Vegetable Center sour: Characterization Utilization, doi: 10.1017/S147926211500057X
East and Southeast Asia, Kamphaeng Saen, Thailand, during (in press).
March and April every year. Plants of the susceptible check va- Duvick DN (2005) The contribution of breeding to yield advances in
riety exhibit 100% susceptibility if planted in the field in this maize (Zea mays L.). Adv Agron 86: 83–145.
period. Based on field screening in this environment, we have FAO (2012) http://faostat.fao.org/site/567/default.aspx#ancor (accessed
developed eight pumpkin lines with field resistance to multiple on 25 March 2016).
unidentified viruses. We are confirming the resistance by artifi- Gogi MD, Ashfaq M, Arif MJ, Sarfraz RM, Nawab NN (2010) Inves-
cial screening against local isolates of CMV, PRSV-W, ZYMV, tigating phenotypic structures and allelochemical compounds of the
and SqLCV. fruits of Momordica charantia L. genotypes as sources of resistance
against Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae).
Crop Prot 29: 884–890.
Cucumber Breeding at AVRDC Hollingsworth R, Vagalo M, Tsatsia F (1977) Biology of melon fly, with
special reference to the Solomon Islands. In: Allwood AJ, Drew RAI
– The World Vegetable Center
(eds) Management of fruit flies in the Pacific. Proceedings of Austra-
lian Country Industrial Agricultural Research 76: 140–144.
Cucumber breeding at AVRDC focused on the development Keneni G, Bekele E, Imtiaz M, Dagne K (2012) Genetic vulnerability of
of predominantly female high yielding tropical lines suitable for modern crop cultivars: Causes, mechanism and remedies. Int J Plant
the Southeast Asian market. We have released one completely Res 2: 69–79.
gynoecious line and five predominantly female lines, which are Krawinkel MB, Keding GB (2006) Bitter gourd (Momordica charantia
also high yielding and remain non-bitter in tropical growing con- L.): a dietary approach to hyperglycaemia. Nutr Rev 64: 331–337.
ditions. Léon J (1994) Mating system and the effect of heterogeneity and het-
erozygosity on phenotypic stability. In: van Ooijen JW, Jansen J (eds)
Biometrics in plant breeding: Applications of molecular markers.

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Proceedings of the 9th meeting of the EUCARPIA section on biomet- Pachner M, Paris HS, Lelley T (2011) Genes for resistance to Zucchini
rics in plant breeding. Wageningen: CPRO-DLO, pp 19–31. yellow mosaic in tropical pumpkin. J Hered 102: 330–335.
Loy BJ (2011) Breeding squash and pumpkins. In: Wang Yi-H, Behe- Rabindranath K, Pillai KS (1986) Control of fruit fly of bitter gourd
ra TK, Kole C (eds.) Genetics, genomics and breeding of cucurbits. using synthetic pyrethroids. Entomon 11: 269–272.
Boca Raton, FL: CRC Press, pp 93–139. WHO 2015. http://www.who.int/diabetes/en/. Accessed 18 Sept 2015.
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al: familiar and exotic cucurbits have Asian origins. HortScience 48: based approach to manage type 2 diabetes and promote health in de-
1078–1089. veloping countries. Wise Folder Hider Pro 4.3.9.199 Crack + License Key {2021} Hort 1106: 161–167.
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ni yellow mosaic virus in Cucurbita moschata. Cucurbit Genet Coop composition of Momordica charantia L. fruits. J Agr Food Chem 39:
Rep 10: 80–81. 1762–1763.

Extreme Picture Finder 3.51.4 Crack - Activators Patch 25
Superfruiter (sf): A New Melon Type with Multiple Small
Seedless Fruits and Increased Yield
Ya’akov Tadmor, Ido Oz, Uzi Sa’ar, Ayala Meir, Fabian Baumkoler,
Nurit Katzir, Efraim Lewinsohn, Vitaly Portnoy, Galil Tzuri,
Arthur A. Schaffer, and Joseph Burger
Center for the Genetic Improvement of Cucurbit Fruit Quality, Agricultural Research Organization,
Newe Ya‘ar Research Center, Ramat Yishay, Israel
e-mail: tadmory@volcani.agri.gov.il

Abstract. We screened an EMS mutation library of melon for unusual fruit and plant phenotypes and identified
a unique recessive mutant that produces more fruit than the wild-type. Fruits were small, one-third of the wild-type
size, and seedless. We called this mutant superfruiter (sf). Indeed, sf decreased fruit size by one-third, but increased
fruit number by five times and on the average increased yield by 67%, in various genetic backgrounds. Thus, sf is
a single recessive gene that significantly breaks yield barriers in melon, without any adverse effect on fruit quality
attributes. We are now developing sf melon varieties by introducing this gene into selected genetic backgrounds.

Introduction a research and breeding tool. Thereafter, thousands of new crop


varieties were released that were bred with new traits obtained
It is widely accepted that organisms are selected according via induced mutation (Ahloowalia et al. 2004, Mba 2013). In ad-
to fitness in their natural habitats. Some of the initial steps in dition, analyses of induced mutants led to the discovery of many
human civilization included the selection of plant phenotypes genes and gene functions by forward and reverse genetics meth-
that had better performance under cultivation. Thus, the organ- ods (Dahmani-Mardas et al. 2010, Bahadur et al. 2015). Induced
ism became selected for its value to humans, or its ability to mutagenesis provides a powerful alternative to natural monogen-
fulfill human’s personal desires, rather than for fitness “in the ic or polygenic variation for identifying functional pathways and
wild” (Pollan 2001). These axioms were the fundamental pillars genes involved in complex traits (Varshney et al. 2014).
of ancient agronomy and horticulture; and traits that were favor- Melon (Cucumis melo L.) is an economically important cu-
able were selected based on their ability to promote the supply curbit crop. Melons exhibit high polymorphism at the DNA level
of food under the conditions in which the growers lived. Adap- that is reflected in their high variation of fruit morphology and
tation to new environments or improvements of these primitive physiology. Melon diversity encompasses fruit size, weight, ex-
cultivars was based on selection of new phenotypes from the ex- ternal and internal color, shape, and aroma. The large within-spe-
isting gene pool. Soon enough, humankind learned, by mixing cies diversity, a small genome size, and a high economic value,
and crossing accessions, to create new variations that served as make melon a convenient and attractive model plant for research.
a new pool from which desired phenotypes were further selected Modern markets of agricultural products are constantly
(Zamir 2001). teamviewer crack reddit index of teamviewer crack - Free Activators looking for novel fruits and vegetables. This is reflected by the
We are therefore constantly looking for methods to increase demand for personal mini fruit; however, decreased fruit size is
variation within a crop. The most common method to increase usually associated with yield reduction. To induce new variation
variation within a crop is to go back to its origins, look for bene- in melon, we conducted chemical mutagenesis on seeds of CEZ,
ficial traits that have been left behind during domestication either a ‘Charentais’-type cantaloupe breeding line. One of the result-
in wild relatives or in primitive varieties and introduce these traits ing M2 families segregated to a unique phenotype easily recog-
Источник: https://www.scribd.com/document/411399236/Conference-Proceedings-Book

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Extreme Picture Finder Crack

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Extreme Picture Finder Crack Download is a powerful application that can be used to find and download digital images, mp3/wmv music and MPG/AVI/WMV videos. Simply enter the website address, select the file to download and save the location, then let Extreme PictureFinder Patch do the rest for you. You can view the downloaded file immediately in the built-in image viewer.

Extreme Picture Finder Full Version Crack has SSD Fresh Licenses key user-friendly interface that allows the rookie to get started with the wizard. You can configure proxy settings and specify a default destination folder. It allows you to choose a site browsing mode between regular gallery and thumbnail gallery. It focuses on websites with links to images on external URLs. Scanning restrictions may include the entire site, the current directory, only the current page or all links. Extreme Picture Finder Serial Key 2021 allows you to use the search function in the downloaded project, view logging details, create a web thumbnail library, and manage file list columns.

As the name suggests, another important feature of the program is the built-in picture finder. Just enter a keyword and the program will find and download thousands of pictures for you. It has different search modes that allow you to change the level of interaction with the download process. With Extreme Picture Finder Registration Key you can let the program find and download only thumbnails, and then see hundreds of images flying towards you within a few seconds-this is the fastest search mode.

You can also free downloadWise Duplicate Finder Pro Crack

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How to Crack Extreme Privazer Crack 4.0.26 With License Key Download 2021 [Latest] Finder 3.51.4 Cracked??

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openSUSE

Last Update: 2021-11-18 19:20 UTC



  • OS Type:Linux
  • Based on: Independent
  • Origin:Germany
  • Architecture:aarch64, armhf, i686, ppc64, ppc64el, s390x, x86_64
  • Desktop:Cinnamon, Enlightenment, GNOME, IceWM, KDE Plasma, LXDE, LXQt, MATE, Xfce
  • Category:Desktop, Server, Live Medium, Raspberry Pi
  • Status:Active
  • Popularity:12 (770 hits per day)
The openSUSE project is a community program sponsored by SUSE Linux and other companies. Promoting the use of Linux everywhere, this program provides free, easy access to openSUSE, a complete Linux distribution. The openSUSE project has three main goals: make openSUSE the easiest Linux for anyone to obtain and the most widely used Linux distribution; leverage open source collaboration to make openSUSE the world's most usable Linux distribution and desktop environment for new and experienced Linux users; dramatically simplify and open the development and packaging processes to make openSUSE the platform of choice for Linux developers and software vendors.



Popularity (hits per day): 12 months: 11 (831), 6 months: 12 (770), 3 months: 12 (720), 4 weeks: 13 (747), 1 week: 15 (783)

Average visitor rating: 8.94/10 from 507review(s).

Источник: https://distrowatch.com/table-mobile.php?distribution=opensuse&pkglist=true&version=tumbleweed

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