Biology of Indian Morels Biology of Indian Morels

T. N. Lakhanpal Onkar Shad Monika Rana

I.K. International Publishing House Pvt. Ltd.

NEW DELHI • BANGALORE Published by I.K. International Publishing House Pvt. Ltd. S-25, Green Park Extension Uphaar Cinema Market New Delhi–110 016 (India) E-mail: [email protected]

ISBN 978-93-80578-24-8

© 2010 I.K. International Publishing House Pvt. Ltd.

All rights reserved. No part of this book may be reproduced or used in any form, electronic or mechanical, including photocopying, recording, or by any information storage and retrieval system, without written permission from the publisher.

Published by Krishan Makhijani for I.K. International Publishing House Pvt. Ltd., S-25, Green Park Extension, Uphaar Cinema Market, New Delhi–110 016. Printed by Rekha Printers Pvt. Ltd., Okhla Industrial Area, Phase II, New Delhi–110 020. Objects in nature have different forms, appearances and names. The truth, however, is that they have a common origin. In this truth of creation, love is the energy that binds all the five elements together. The entire creation is suffused with the Divine energy of love.

To the Divine Lotees Feet of Sri Sathya Sai Baba

&

To my guru and mentor Shri K.G. Mukerjee Foreword

I am happy to write a forward for the monography ‘Biology of Indian Morels’ penned by Prof. T.N. Lakhanpal and his associates, Dr. O. S. Shad and Dr. Monika Rana. Morels have attracted the poor and the rich, the young and the old alike since times immemorial all over the world because of their taste and excellent flavour. Called ‘Guchhi’ in the Indian market morels are one of the most important commercial wild edible fungi. The research on the morels has been primarily centered on taxonomy and on the efforts for domestication. All the biological aspects, however, are perhaps being covered for the first time in this monograph. Morels in India grow mainly in the Himalayan region that is why most of the research work has come from researchers located in and around the Himalayan region especially Jammu and Kashmir and Himachal Pradesh. Though different aspects have now been worked our, the artificial cultivation still eludes the researchers and that is the reason for so high cost presently. The authors of this monograph have investigated during the last three decades and have for the first time attempted to compile and consolidate the available literature along with their own work into this voluminous monograph. There are fourteen chapters in this monograph each chapter is devoted to one aspect of morel biology. The monography begins with the overview provides information on life cycle pattern, given classical concept of morel taxonomy followed by molecular characterization of the species. Subsequently, it provides information on various ecological, physiological, rhizomorphospheric aspects and associations, cytology, mycological, sociobiological, nutraceutical potential and culinary aspects. Information collected from the local people and the impact of morel collection on the economy of the people has been discussed in detail. The data on medicinal uses, collection, drying and trading, nomenclatures, colour variation, relationship to snow fall and forest types reveals many startling facts about these small viii Foreword organisms with big potential. In nutshell this monograph provides excellent original information on almost all aspect of the morel biology. May be this compilation inspires researcher and one day the technology for its cultivation is developed and perfected. That will be the golden day in the history of the fungal biology and biodiversity. I congratulate Prof. T. N. Lakhanpal, Dr. O. S. Shad and Dr. Monika Rana for sincere efforts and hard work in preparing the Monograph “Biology and Indian Morel” in a highly scientific and qualitative way and the monograph will be highly useful to Teachers, Scientists, Scholars and Students. I am sure the monograph will find ample use in all the libraries and research institutes.

Prof. S. Kannaiyan 2nd Proof 12/02/08

Preface

Morels have been ‘moral’ booster mushrooms as they have inspired mycologists to keep research on them alive so as to achieve their artificial cultivation. Morels are king mushrooms and have attracted human attention since time immemorial. They have earned appreciation for their excellent flavour and culinary properties. Technically morels belong to the genus Morchella and are commonly called ‘Guchhi’ in the Indian market. They are presently the costliest mushrooms selling at Rs. 7000-8000 per kg in the Indian market. The senior author was fascinated by the morels as a postgraduate student in the late sixties at the Botany Department, Punjab University, Chandigarh. He was exposed first to the colourful world of fungi when he joined fungal forays to the N-W Himalayas with groups led by late Prof. K.S. Thind. The interest was revived in the beginning of 1980s after he joined Department of Biosciences H P University Shimla in 1976. Systematic work on morels was initiated and executed from 1980 onwards with great zeal. This progressed further with the enthusiasm of the second author who worked for his M.Phil. (1985) and then Ph.D. (1989) on classical aspects followed by the third author who completed molecular characterization of the species for her Ph.D. in 2005. In between some interludes on ethnomycology were provided by Mangala Gowry, Meenakshi Rathod and Mitu Sharma. Hence, this monograph on ‘Biology of Indian Morels’ is an outcome of studies initiated in 1980 and carried through all the years till 2005 steadily but continuously and earnestly. During all these years almost all aspects of morel biology have been investigated. However, the cultivation of morels still eludes the scientists the world over. x Preface

The monograph has a number of chapters, each devoted to one aspect of study. It begins with an overview of morel biology, the life cycle pattern, the classical and molecular systematic treatment, the ecological and physiological aspects, the nutritional and nutraceutical components, mycorrhizal and sociobiological and ethnomycological investigations, cytological studies and the traditional and modern recipes for morel consumptions. All the chapters are documented with tables, figures and original photographs. It is hoped this monograph will interest botanists and in particular mycologists the world over. Suggestions for improvement are welcome.

T.N. Lakhanpal Onkar Shad Monika Rana Acknowledgements

Many individuals, directly or indirectly, have helped in the accomplishment of this endeavour and mere acknowledgement of their help in a few formal words may not actually be truly representative of the assistance rendered. First and the foremost acknowledgement are to the Almighty for enabling us to undertake the task and successfully accomplishing it. Without His grace and unseen hand, nothing would have been possible. Our grateful thanks to Prof. K.S. Thind, Panjab University, Chandigarh, who inspired the senior author to undertake this type of work, to Prof. K.G. Mukerji. University of Dehli for the role of a friend, philosopher and guide in the execution and publishing of this work. Thanks in no less measure to Professors C.V. Subramanian, R.L. Munjal, J.N. Kapoor, T.N. Kaul, K. Natrajan, R.P. Purkayastha, Dr. A.V. Sathe, for help in various ways. Thanks are also due to Dr. Roy Watling, Edinburgh, Dr. Gary Lincoff, U.S.A., Prof. James Trappe, Michael Castillano, David Arora, Randy Molina, Nancy Weber Smith, Oregon, Corvallis, U.S.A., Prof. E. Horak, Switzerland and Prof. T. Hongo, Japan. The senior author is thankful to all his students and colleagues who were ever ready for rendering help on all occasions. Grateful thanks are also due to Prof. H.S. Garcha, PAU Ludhiana, Dr. R.P. Tiwari, Director, Dr. S.R. Sharma, NMRC, Solan, Prof. C.L. Jandik, Solan, Prof. Dalip Malhotra, Prof. S.K. Sood, Prof. V.K. Mattu, Dr. K.S. Kapoor, and Dr. Vineet Jistu for moral support and encouragement. Help extended by Dr. S.K. Chakrabarti CPRI Shimla during the studies on molecular aspect is gratefully acknowledged. Our grateful thanks to our parents and family members for their understanding and encouragement. xii Acknowledgements

Financial help from the Department of Science and Technology, and Dept. of Biotechnology, GOI, New Delhi to the senior author is duly acknowledged. This grant made possible the execution and the completion of the work.

T.N. Lakhanpal Onkar Shad Monika Rana Contents

Foreword vii Preface viii Acknowledgements xi 1. Biology of Morels–An Overview 1 Introduction 2 Habit and Habitat 2 Systematic Treatment 2 Nutritive Value 4 Ecological Aspects 6 Rhizomorphosphere Associations 8 Ethnomycological Significance 8 Production and Trade 9 2.The Life Cycle Pattern 11 The Ascocarp 12 Ascospores and Mycelium 12 Heterokaryon Formation 13 Sclerotial Development 15 Ascocarp Development 18 Cultivation Aspects 18 xiv Contents

3.Systematic Treatment of Morels–I 21 Introduction 22 Review of Literature 22 Review of Work done in India 24 Systematic Treatment of Species of Morchella 25 Key to the Indian Species of Morchella 25 Discussion 35 4.Systematic Treatment of Morels–II: Genotypic Polymorphism and Diversity 39 Introduction and Review of Literature 40 Observations 45 Similarity Analysis and Clustering of Data 49 Discussion 51 5. Ecological Aspects 53 Introduction and Review of Literature 54 Observations and Discussion 56 6. Physiological Aspects of Morels65 (I) General Physiological Aspects 66 Introduction 66 Review of Indian Literature 67 Observations and Discussion 68 (II) Analysis of Enzymatic Potential 71 Introduction 71 Review of Literature 79 Observations and Discussion 80 7.Nutritive Composition 83 Introduction 84 Review of Literature 84 Observations and Discussion 87 8.Nutraceutical Potential 91 Introduction and Review of Literature 92 Milestones in Nutraceutical Research 95 Observations and Discussion 98 9. Rhizomorphosphere Relationships 109 Introduction and Review of Literature 110 Observations and Discussion 110 10. Ethnomycological Aspects 121 Introduction and Review of Literature 122 Observations and Discussion 125 Contents xv

11.Sociobiological Aspects 143 Introduction 144 The People 144 Agriculture 144 Agro-Climatic Conditions 145 Horticulture 146 Animal Husbandry 148 Land Use Pattern 148 Production and Trade of Morels 149 Socio-Economic Profile of Morel Collecting Population 150 12.Mycorrhizal Association of Morels 153 Introduction and Review of Literature 154 Observations and Discussion 155 13. Cytology of Ascus Development in Morels 159 Introduction 160 Review of Literature 160 Collection and Fixation of Material 161 Observation and Discussion 161 14. Culinary Aspects of Morels 167 Introduction 168 Review of Literature 169 Observations 171 Annexures 185 References 217 Index 243 Abbreviations and Symbols

" — inches % — per cent / — per & — and mg — microgram ml — microlitre mm — micrometre $ — Dollar AD — Anno Domini amsl — above mean sea level BC — Before Christ BMR — Basic Metabolic Rate bp — base pairs c.f. — cited from cms — centimetres conc. — concentrated d.w. — dry weight Dia. — diameter dil. — dilute DNA — Deoxyribonucleic Acid Abbreviations and Symbols xvii

e.g. — for example ed. — edition Ed. — editor Eds. — editors et al., — et. alia; and others etc. — et cetera Fig. — Figure g — gram H.P. — Himachal Pradesh

H2SO4 — Sulphuric acid HCL — Hydrocholoric acid HDP — Health Defence Potentiators HPUB — herbarium, Department of Biosciences, H.P. University, Shimla hrs. — hours i.e. — that is ICBN — International Code of Botanical Nomenclature J&K — Jammu and Kashmir Kcal — Kilo calories Kg — Kilogram l — litre l.c. — Loco citato: at the place cited m — metre mg — milligram mgs — miligrams min — munutes ml — millilitre msl — mean sea level N.W. — north West ng — nano gram °C — Degree Celsius PCR — Polymerase Chain Reaction PDA — Potato Dextrose AGar ppm — parts per millon RAPD — Random Amplified Polymorphic DNA RFLP — Restriction Fragment Length Polymorphism rpm. — rate per minute sp. — Speces (Singular) spp. — Species (Plural) xviii Abbreviations and Symbols

sq km — Square kilometre sq m — Square metre Syn. — Synonym TDF — Total dietary fibre U.P — Uatter Pradesh var. — variety vern. — vernacular viz. — videlicet; namely vol.(s) — volume(s) v/v — volume/volume w.r.t — with respect to CHAPTER Biology of Morels–An 1 Overview

CHAPTER OUTLINE F Introduction

F Habit and Habitat

F Systematic Treatment

F Nutritive Value

F Ecological Aspects

F Rhizomorphosphere Associations

F Ethnomycological Significance

F Production and Trade

Chap-01.p65 1 6/28/10, 10:08 AM 2 Biology of Indian Morels

INTRODUCTION

The morels comprising the genus Morchella, are commonly referred to as ‘Guchhi’ in the Indian Market. At present, it is the costliest and most sought after edible fungus in the world. The morels are highly appreciated for their culinary aspects and gastronomical delights and excellent flavour. Even the mycelia are known to possess the same nutritive value as the sporophores (fruit body). The fructifications constitute the item of commerce and are exported from India to Europe and United States of America.

HABIT AND HABITAT

Species of Morchella in India occur primarily in N-W Himalaya, though there are some reports from sub-hilly areas and plains as well. The species grow on soil rich in organic matter, in loamy soil, in pure and mixed coniferous forests, broad leaved forests, in open areas or grasslands, in apple orchards or under various types of shrubs in the open under direct and indirect shade. Morels mainly appear during the spring season from March to May when snow begins to melt and is accompanied by some precipitation. Species of Morchella have, however, been recorded/collected during the rainy season as well (Wakode, 1983; Lakhanpal and Shad, 1986 a, b; Shad, 1989, Kamal, 2005 and Kaviyasaran et al., 2006), in the months of September and October, repeatedly from some parts of India and various parts of Himachal Pradesh including lower hills which do not experience snowfall (Lakhanpal and Shad, 1986 a, b). Morchella fruiting bodies have also been reported from paddy field bunds (in Jammu) and maize fields (in Solan) (Jandaik and Sharma, 1995). Majority of the Morchella species have been collected from N-W Himalayan region, especially Jammu & Kashmir (Cooke, 1870; Sydow and Butler, 1911; Ghosh and Pathak 1962; Batra and Batra, 1963; Waraitch, 1976 and Kaul, 1976, 1981), Himachal Pradesh; (Sohi et al., 1965; Lakhanpal and Shad, 1986) and Uttar Pradesh (Hennings, 1901 and Theissen, 1911). Occasional reports about collection of Morchella vulgaris Boud. from Assam forests, M. deliciosa from Amritsar (Punjab) and M. conica Pers. from central India have also been given by Bhattacharya and Baruah (1953), Purkayastha and Chandra (1985), and Ghurde and Wakode (1981), respectively. The validity of M. vulgaris from Assam forests, however, has been doubted by Ghosh and Pathak (1962). The distribution/collection of different Morchella species in India is given in Table 1.1.

SYSTEMATIC TREATMENT

The genus Morchella is one of the oldest genera of Pezizales. It was established as early as 1719 by Dillenius. It has undergone little change till today regarding its limits. Though at times Mitrophora Lav. has been included in Morchella (Saccardo, 1889; Seaver, 1928 and Rifai, 1968), but majority of the researchers prefer to treat it as a separate genus, distinguishable by a pileus free from stipe in the lower region (Dennis, 1960, 1968 and Moser, 1963). Opinions have differed with regard to the circumscription of the species in the genus Morchella, so that variable number of species have been recognized from time to time. Groves and Hoare (1953) and Dennis (1960, 1968) recognized three species, whereas Seaver (1928)

Chap-01.p65 2 6/28/10, 10:08 AM Biology of Morels–An Overview 3

Table 1.1 Distribution of Morchella species in India

Species State Reference(s) M. angusticeps Peck. HP Sohi et al. (1965); Munjal et al. (1977); Jandaik et al. (1978); Lakhanpal and Shad (1986) Kashmir Ghosh and Pathak (1962); Batra and Batra (1963); Kaul (1976); Waraitch (1976) Maharashtra Wakode (1963) M. conica Pers. HP Sohi et al. (1965); Munjal et al. (1977); Lakhanpal and Shad (1986) J & K Batra and Batra (1963); Waraitch (1976); Kaul (1976) UP Hennings (1901) Maharashtra Ghurde and Wakode (1981) M. crassipes (Vent.) Pers. ex Fr. HP Lakhanpal and Shad (1986) Kashmir Waraitch (1976) M. deliciosa Fr. HP Munjal et al. (1977); Lakhanpal and Shad (1986) Kashmir Ghosh and Pathak (1962); Batra and Batra (1963); Kaul (1976); Waraitch (1976) Maharashtra Wakode (1983) UP Ghosh and Pathak (1962) M. esculenta (L.) Pers. ex Fr. HP Sohi et al. (1965); Lakhanpal and Shad (1986) Kashmir Ghosh and Pathak (1962); Batra and Batra (1963); Waraitch (1976); Kaul (1976) Maharashtra Wakode (1983) UP Ghosh and Pathak (1962); Sydow and Butler (1911)

Western Ghats Kaviyasaran et al. (2006) M. semilibera DC. ex Fr. (Syn.) HP Sohi et al. (1965); Lakhanpal and Shad (1986) Kashmir Ghosh and Pathak (1962); Kaul (1976)

recognized six species and Boudier (1907) as many as thirty-two species. The delimitation depended upon the weightage one gives to the morphological variations. British mycologists follow a conservative concept and recognize six species, whereas American mycologists tend to split morels into a number of species and recognize more than six species. The earliest record of Morchella from India is by Cooke (1870) as M. gigasporsa Cooke, but this species is now a synonym of M. bohemica Krombh (see Butler and Bisby, 1931) and presently it is placed in the genus Verpa. All the six species considered as valid in the genus Morchella have been recorded from India. These are M. deliciosa Fr. (delicious morels), M. esculenta (L) Pers. ex Fr. (common morel), M. conica Pers. (conical morels), M. crassipes (thick stemmed morel), M. angusticeps Peck. (black morel) and M. hybrida (hybrid morel). The correct name for M. hybrida is M. semilibera DC. ex Fr. The former is untenable (Rifai, 1968) as it was not accepted by Fries (1822). The report of M. vulgaris Boud., which is a synonym of M. esculenta (L.) Pers. ex Fr.

Chap-01.p65 3 6/28/10, 10:08 AM Biology Of Indian Morels

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