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NEW AND FUTURE DEVELOPMENTS IN
MICROBIAL BIOTECHNOLOGY AND
BIOENGINEERING
NEW AND FUTURE
DEVELOPMENTS IN
MICROBIAL
BIOTECHNOLOGY
AND
BIOENGINEERING
Sustainable Agriculture:
Microorganisms as Biostimulants
Edited by

Harikesh Bahadur Singh

Department of Biotechnology, GLA University, Mathura, India

Anukool Vaishnav
Department of Biotechnology, GLA University, Mathura, India; Agroecology and Environment,
Agroscope, Zürich, Switzerland
Elsevier
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 Contents

Contributors ix 3. Bacillus rhizobacteria: A versatile

About the Editors xiii
Preface xv
biostimulant for sustainable agriculture
S.R. Prabhukarthikeyan, U. Keerthana, Mathew S Baite,
P. Panneerselvam, Debasis Mitra, R. Naveen Kumar,
1. Role of microorganism as new C. Parameswaran, B. Cayalvizhi, A. Muthu Kumar,
generation plant bio-stimulants: S. Harish, P.C. Rath

An assessment 3.1 Introduction 33

Deepali Shukla, Piyush Shukla, Ashmita Tandon, 3.2 Diversity of Bacillus species 34
Poonam C. Singh, Jayandra Kumar Johri
3.3 Direct mechanism of plant growth
promotion 35
1.1 Background 1
3.4 Indirect mechanism 37
1.2 Introduction of plant bio-stimulants 2
3.5 Future prospects 40
1.3 Basic mechanism of bio-stimulants 2
References 40
1.4 Sources of plant bio-stimulants 2
1.5 Microbes as plant bio-stimulant 3 4. Arbuscular mycorrhizae, a treasured
1.6 Role of microbes in nutrient uptake/
stimulation 8
symbiont to agriculture
1.7 Conclusions 9 Ajay Nair, Archana S. Rao, L. Bhanu, Veena S. More,
K.S. Anantharaju, Sunil S. More
References 10
4.1 Introduction to mycorrhiza 45
2. Exploiting biostimulant properties 4.2 VAM in agriculture 48
of Trichoderma for sustainable 4.3 Application of AMF in bioremediation 55
plant production 4.4 Renaturation and afforestation 56
4.5 Mass production of VAM: the past, present
Ramón Pelagio-Flores, Saraí Esparza-Reynoso, Jesús Salvador
López-Bucio, José López-Bucio and future 57
4.6 Conclusion 59
2.1 Introduction 17 References 59
2.2 Trichoderma metabolism: from decomposers to
plant growth promoters 19 5. Micro and macroalgae: A potential
2.3 Trichoderma-plant chemical dialogue 19 biostimulant for abiotic stress management
2.4 Trichoderma-induced resistance to plant and crop production
pathogens 20 P. Kiruthika Lakshmi, S. Meenakshi
2.5 Trichoderma and plant nutrition 22
2.6 Soil acidification in Trichoderma-plant 5.1 Introduction 63
interactions 25 5.2 Review of literature and recent developments 64
2.7 Salt stress tolerance mediated by 5.3 Conclusion and future prospects 76
Trichoderma 25 References 77
2.8 Conclusions and future prospects 26
References 27

v
vi CONTENTS

6. Fluorescent pseudomonads: 8.5 Genetics involved in nutrient cycling 121

A multifaceted biocontrol agent for 8.6 Conclusions 124
References 126
sustainable agriculture
U. Keerthana, S.R. Prabhukarthikeyan, Mathew S. Baite, 9. Role of soil microbes in
Manoj K. Yadav, R. Naveen Kumar, A. Muthu Kumar, S. Raghu,
S. Aravindan, C.P. Rath
micronutrient solubilization
Bisweswar Gorain, Srijita Paul, Manoj Parihar
6.1 Introduction 88
6.2 Species diversity of Fluorescent 9.1 Introduction 131
Pseudomanads 83 9.2 Importance of micronutrients in plant
6.3 Mechanisms of Fluorescent Pseudomanads 84 nutrition 132
6.4 Future prospects 88 9.3 Sources and pools of micronutrients in soil and
References 89 their significance in plant uptake 133
9.4 Factors affecting the availability of
7. Role of Piriformospora indica in inducing micronutrients 133
9.5 Influence of rhizosphere in micronutrient
soil microbial communities and drought availability 134
stress tolerance in plants 9.6 Soil pH and pE as an indicator of micronutrient
Suryansh Rajput, Preeti Sengupta, Isha Kohli, Ajit Varma, availability 134
Prashant Kumar Singh, Naveen Chandra Joshi 9.7 Micronutrients 135
9.8 Conclusion and future perspectives 145
7.1 Introduction 93
References 146
7.2 Soil microbial communities: benign hidden
players in plant growth 94 10. Sustainable induction of systemic
7.3 P. indica: an overview 95 resistance in response to potential
7.4 Basic mechanisms in plants to counter drought
stress 101
biological control agents in crops
7.5 Morphological and physiological innate Ali Anwar, Efath Shahnaz
responses in plants against drought stress 102
10.1 Introduction 151
7.6 Multidimensional contribution of P. indica in
10.2 Novel scenario of biological control 153
providing tolerance against drought stress 103
10.3 Suppressive soils pathogens 154
7.7 P. indica mediated adaptative responses generated
10.4 Potential in PGPR 155
in rice plants to cope up drought stress 105
10.5 Induction of systemic resistance 157
7.8 Scope of P. indica for the promotion of
10.6 Fungal BCAs 166
sustainable agriculture in xerophytic
10.7 Potental of non-pathogenic strains 167
habitats 106
10.8 Conclusion and future prospects 168
7.9 Conclusion 107
References 169
References 107
11. Psychrophilic microbes: Biodiversity,
8. Microbes-based bio-stimulants towards beneficial role and improvement of
sustainable oilseeds production: Nutrients cold stress in crop plants
recycling and genetics involved Sumit Kumar, Bana Sravani, Tulasi Korra, Lopamudra Behera,
A.L. Singh, Kirti Rani, Rupak Jena, Praveen Kona, Diptanu Datta, Pushpendra Kumar Dhakad, M.K. Yadav
Kiran K. Reddy, K. Gangadhara
11.1 Introduction 177
8.1 Introduction 111 11.2 Historical background 180
8.2 Soil microbes and plant interactions 112 11.3 Biodiversity of psychrophilic microbes 180
8.3 Geochemical changes in plant rhizosphere and 11.4 Mechanisms of adaptation of psychrophilic
release of mineral nutrients 115 microbes 182
8.4 VAM fungi for efficient nutrient acquisition and 11.5 Psychrophilic microbes used in crop
mobilization 119 improvement 186
 CONTENTS
11.6 The beneficial role of psychrophilic microbes in
crop performance 189
11.7 Conclusion and future prospects 192
Reference 193
12. Role of plant-associated bacteria as
bio-stimulants in alleviation of
chromium toxicity in plants
Pratishtha Gupta, Vipin Kumar, Rupa Rani
12.1 Cr toxicity to the environment 199
12.2 Strategies of Cr remediation from contaminated
environment 200
12.3 Plant growth promoting rhizobacteria and their
beneficial traits 200
12.4 Cr induced oxidative stress in plants and
anti-oxidative enzymes 203
12.5 PGPR and phytoremediation 205
12.6 Case study of Cr phytoremediation mediated by
root-associated bacteria 207
12.7 Conclusion 208
References 209
13. Microbe-based plant biostimulants and
their formulations for growth promotion
and stress tolerance in plants
Abhik Mojumdar, Himadri Tanaya Behera,
Smrutiranjan Das, Lopamudra Ray
13.1 Introduction 213
13.2 Microbes as plant biostimulants 215
13.3 Mechanism of development of microbe-based
plant biostimulants 217
13.4 Microbial bioformulation based plant
biostimulants 218
13.5 Microbes as biofertilizers 220
13.6 Biopesticides 222
13.7 Significance of microbes in abiotic and biotic
stress alleviation 223
13.8 Challenges and future prospects 224
13.9 Conclusions 225
References 225
14. Microbial consortia for augmentation
of plant growth–revisiting the promising
approach towards sustainable agriculture
Niharika Ghoghari, Krishna Bharwad, Akshita Champaneria,
Shalini Rajkumar
14.1 Rhizosphere: a nutrient rich niche 234
14.2 Microbial marketing strategies 234
vii
14.3 Plant microbe interactions 234
14.4 Microbe-microbe interactions 236
14.5 Plant probiotics 237
14.6 Plant growth promoting rhizobacteria
(PGPR) 237
14.7 Nitrogen fixation 237
14.8 Mineral acquisition 238
14.9 Phytohormone production 238
14.10 Prevention of diseases and development
of ISR 238
14.11 Biocontrol agents 238
14.12 Biostimulants 239
14.13 Microbial consortia: the dynamics of
co-operation 240
14.14 Binary consortium 243
14.15 Three or multi partner consortium
development 244
14.16 Multi-omics for development of microbial
consortia for plant growth promotion 248
References 250
15. Phosphate solubilization
by microorganisms
Neimar de Freitas Duarte, Christiane Abreu Oliveira Paiva,
Marcela Claudia Pagano, Eduardo Jose Azevedo Correa
15.1 Introduction 257
15.2 Research the selection of phosphate-solubilizing
microbes 268
15.3 Bioinoculants containing strains of P solubilizing
microorganisms and biomaphos - an example of
a successful case in Brazil 269
References 274
16. Fungal endophytes as biostimulants of
secondary metabolism in plants: a
sustainable agricultural practice for
medicinal crops
Tuyelee Das, Abhijit Dey, Devendra Kumar Pandey,
Joginder Singh Panwar, Samapika Nandy
16.1 Introduction 283
16.2 Why do we need to study fungal-medicinal
plant interaction to make secondary
metabolites? 285
16.3 Role of endophytic fungi in production of
secondary metabolites; host-endophyte
relationship 286
16.4 Metabolic interactions of plant
endophytes 299
viii CONTENTS

16.5 Different strategies to exploit fungal endophytes 18.6 Impact of microbiome-based approaches on the

as biostimulants for production of commercially health of plant and soil 338
important plant-derived compounds 301 18.7 Future of microbiome-based approaches in
16.6 Secondary metabolic compounds produced by enhancing soil health: integration of
medicinal plants endophytic fungi in vitro 304 metagenomics and metabolomics approaches
16.7 Conclusion 308 with designing of synthetic communities 339
Acknowledgment 308 18.8 Conclusion 341
References 308 Acknowledgement 342
References 342
17. Plant growth promoting rhizobacteria
from the perspectives of tea 19. Deciphering microbial consortium
plantations and diseases from termite gut for biofertilizer
Stevenson Thabah, S.R. Joshi consortium formulation
Sujit Ghosh, Ambarish S. Vidyarthi, Vijay Kumar, Priyanka Jha
17.1 Introduction 315
17.2 Tea cultivation in India 316 19.1 Introduction 345
17.3 Tea varieties 316 19.2 Material and methods 346
17.4 Shade trees in tea plantations 317 19.3 Results and discussions 347
17.5 Pests and diseases of tea 318 19.4 Conclusion 351
17.6 Tea rhizosphere 319 Acknowledgements 351
17.7 Rhizospheric activity 319 References 351
17.8 Plant growth promoting rhizobacteria
(PGPR) 321 20. Revivification of rhizobacteria-
17.9 PGPR and prospective benefits to tea promoting plant growth for sustainable
plants 322
agricultural development
17.10 PGPR as biocontrol agents in tea
Zahra Dehghanian, Khashayar Habibi, Monire Mones Sardrodi,
cultivation 326
Khosro Balilashaki, Behanm Asgari Lajayer, Tess Astatkie
17.11 Tea plantations and microbial
colonization 326 20.1 Introduction 353
17.12 Conclusion 328 20.2 Rhizosphere soil 354
References 329 20.3 Plant growth promoting rhizobacteria
(PGPR) 354
18. Microbiome-based approaches to 20.4 PGPR in farming 354
enhance soil health in arable land 20.5 Bio-fertilization 355
Shivani Khatri, Shubham Dubey, Shilpi Sharma 20.6 The PGPR biological control agents 355
20.7 Mechanisms of direct 356
18.1 Introduction 333 20.8 Indirect mechanisms 359
18.2 Conventional microbe-based approach for 20.9 Sustainability of agriculture and future
enhancement of soil health 334 perspective 362
18.3 Limitations associated with conventional 20.10 Conclusions 362
approaches 335 References 363
18.4 Microbiome: a brief overview 335
18.5 Approaches used to engineer the Index 369
microbiome 336
 Contributors
K.S. Anantharaju Department of Chemistry, Day-
ananda Sagar Collge of Engineering, Bangalore,
India
Ali Anwar Division of Plant Pathology, Faculty
of Horticulture, SKUAST-Kashmir, Srinagar,
J&K, India
S. Aravindan ICAR-National Rice Research In-
stitute, Cuttack, Odisha, India
Tess Astatkie Faculty of Agriculture, Dalhousie
University, Truro, Canada
Mathew S Baite ICAR-National Rice Research
Institute, Cuttack, Odisha, India
Khosro Balilashaki Department of Horticultur-
al Science, Faculty of Agriculture, University of
Guilan, Guilan, Iran
Himadri Tanaya Behera School of Biotechnol-
ogy, Kalinga Institute of Industrial Technology
(KIIT), Bhubaneswar, Odisha, India
Lopamudra Behera Department of Mycology
and Plant pathology, Institute of Agricultural
Sciences, Banaras Hindu University, Varanasi,
India
L. Bhanu School of Basic and Applied Sciences,
Dayananda Sagar University, Bangalore, India
Krishna Bharwad Institute of Science, Nirma
University, Sarkhej-Gandhinagar Highway,
Ahmedabad Gujarat, India
B. Cayalvizhi ICAR-National Rice Research In-
stitute, Cuttack, Odisha, India
Akshita Champaneria Institute of Science, Nir-
ma University, Sarkhej-Gandhinagar Highway,
Ahmedabad, Gujarat, India
Eduardo Jose Azevedo Correa Agricultural Re-
search Company of Minas Gerais (EPAMIG),
Pitangui, MG
Smrutiranjan Das School of Biotechnology, Ka-
linga Institute of Industrial Technology (KIIT),
Bhubaneswar, Odisha, India
ix
Tuyelee Das Department of Life Sciences, Presi-
dency University, Kolkata, India
Diptanu Datta Department of Plant Pathol-
ogy, Collage of agriculture, Odisha University
of Agriculture and Technology, Bhubaneswar,
Odisha, India
Zahra Dehghanian Department of Biotechnol-
ogy, Faculty of Agriculture, Azarbaijan Shahid
Madani University, Tabriz, Iran
Abhijit Dey Department of Life Sciences, Presi-
dency University, Kolkata, India
Pushpendra Kumar Dhakad Holy Cross Kri-
shi Vigyan Kendra, Canary Hill, Hazaribag,
Jharkhand, India
Neimar de Freitas Duarte Minas Gerais Federal
Institute of Education, Science and Technology
(IFMG), Brazil
Shubham Dubey Department of Biochemical
Engineering and Biotechnology, Indian Insti-
tute of Technology Delhi, Hauz Khas, New
Delhi, India
Saraí Esparza-Reynoso Instituto de Investiga-
ciones Químico-Biológicas, Universidad Mi-
choacana de San Nicolás de Hidalgo, Edificio
B3, Ciudad Universitaria, Morelia, Michoacán,
México
K. Gangadhar ICAR-Directorate of Groundnut
Research, Junagadh, Gujarat, India
Niharika Ghoghari Institute of Science, Nirma
University, Sarkhej-Gandhinagar Highway,
Ahmedabad, Gujarat, India
Sujit Ghosh Department of Botany, J.K. Col-
lege, Purulia, West Bengal, India
Bisweswar Gorain ICAR-Central Soil Salinity
Research Institute RRS Bharuch, Gujarat, India
x CONTRIBUTORS
Pratishtha Gupta Applied Microbiology Labo-
ratory, Department of Environmental Science
and Engineering, Indian Institute of Technol-
ogy (ISM), Dhanbad, Jharkhand, India
Khashayar Habibi Department of Plant Breed-
ing and Biotechnology, Faculty of Agriculture,
University of Tabriz, Tabriz, Iran
S. Harish Tamil Nadu Agricultural University,
Coimbatore, Tamil Nadu, India
Rupak Jena ICAR-Directorate of Groundnut
Research, Junagadh, Gujarat, India
Priyanka Jha Amity Institute of Biotechnology,
Amity University, Major Arterial Road, Action
Area II, Kolkata, West Bengal, India
Jayandra Kumar Johri Department of Plant
Microbe Interaction, CSIR-NBRI, Lucknow, UP,
India
Naveen Chandra Joshi Amity Institute of Mi-
crobial Technology, Amity University, Noida,
Uttar Pradesh, India.
S.R. Joshi Microbiology Laboratory, De-
partment of Biotechnology & Bioinformat-
ics, North-Eastern Hill University, Shillong,
Meghalaya, India
U. Keerthana ICAR-National Rice Research In-
stitute, Cuttack, Odisha, India
Shivani Khatri Department of Biochemical En-
gineering and Biotechnology, Indian Institute
of Technology Delhi, Hauz Khas, New Delhi,
India
Isha Kohli Amity Institute of Microbial Technol-
ogy, Amity University, Noida, Uttar Pradesh,
India
Praveen Kona ICAR-Directorate of Groundnut
Research, Junagadh, Gujarat, India
Tulasi Korra Department of Mycology and
Plant pathology, Institute of Agricultural Sci-
ences, Banaras Hindu University, Varanasi,
India
Sumit Kumar Department of Mycology and
Plant pathology, Institute of Agricultural Sci-
ences, Banaras Hindu University, Varanasi, India
Vipin Kumar Applied Microbiology Laboratory,
Department of Environmental Science and
Engineering, Indian Institute of Technology
(ISM), Dhanbad, Jharkhand, India
Vijay Kumar Department of Biotechnology,
Lovely Faculty of Technology and Sciences,
Lovely Professional University, Phagwara,
Punjab, India
Behanm Asgari Lajayer Health and Environ-
ment Research Center, Tabriz University of
Medical Sciences, Tabriz, Iran
P. Kiruthika Lakshmi Department of Micro-
biology, The Madura College, Madurai, Tamil
Nadu, India
Jesús Salvador López-Bucio CONACYT, Insti-
tuto de Investigaciones Químico-Biológicas,
Universidad Michoacana de San Nicolás de Hi-
dalgo. Edificio B3, Ciudad Universitaria, More-
lia, Michoacán, México
José López-Bucio Instituto de Investigaciones
Químico-Biológicas, Universidad Michoacana
de San Nicolás de Hidalgo, Edificio B3, Ciudad
Universitaria, Morelia, Michoacán, México
S. Meenakshi VMJ School, Madurai, Tamil
Nadu, India
Debasis Mitra ICAR-National Rice Research In-
stitute, Cuttack, Odisha, India
Abhik Mojumdar School of Biotechnology, Ka-
linga Institute of Industrial Technology (KIIT),
Bhubaneswar, Odisha, India
Sunil S. More School of Basic and Applied Sci-
ences, Dayananda Sagar University, Bangalore,
India
Veena S. More Department of Biotechnology,
Sapthagiri College of Engineering, Bangalore,
India
A. Muthu Kumar Faculty of Agriculture, An-
namalai University, AnnamalaiNagar, Tamil
Nadu, India
Ajay Nair School of Basic and Applied Scienc-
es, Dayananda Sagar University, Bangalore,
India
Samapika Nandy Department of Life Sciences,
Presidency University, Kolkata, India
R. Naveen Kumar ICAR-National Rice Re-
search Institute, Cuttack, Odisha, India
 CONTRIBUTORS
Christiane Abreu Oliveira Paiva Brazilian Ag-
ricultural Research Corporation (EMBRAPA),
Sete Lagoas, Minas Gerais, Brazil
Marcela Claudia Pagano Federal University of
Minas Gerais, Belo Horizonte, Minas, Gerais,
Brazil
Devendra Kumar Pandey Department of Bio-
technology, Faculty of Technology and Scienc-
es, Lovely Professional University, Phagwara,
Punjab, India
P. Panneerselvam ICAR-National Rice Research
Institute, Cuttack, Odisha, India
Joginder Singh Panwar Department of Biotech-
nology, Faculty of Technology and Sciences,
Lovely Professional University, Phagwara,
Punjab, India
C. Parameswaran ICAR-National Rice Research
Institute, Cuttack, Odisha, India
Manoj Parihar ICAR-VPKAS, Almora Uttara-
khand, India
Srijita Paul ATC and SAMETI, Ramakrishna
Mission Ashrama, Kolkata, W.B., India
Ramón Pelagio-Flores Facultad de Químico
Farmacobiología, Universidad Michoacana de
San Nicolás de Hidalgo, Morelia, Michoacán,
México
S.R. Prabhukarthikeyan ICAR-National Rice
Research Institute, Cuttack, Odisha, India
S. Raghu ICAR-National Rice Research Insti-
tute, Cuttack, Odisha, India
Shalini Rajkumar Institute of Science, Nirma
University, Sarkhej-Gandhinagar Highway,
Ahmedabad, Gujarat, India
Suryansh Rajput Amity Institute of Microbial
Technology, Amity University, Noida, Uttar
Pradesh, India
Kirti Rani ICAR-Directorate of Groundnut Re-
search, Junagadh, Gujarat, India
Rupa Rani Applied Microbiology Laboratory,
Department of Environmental Science and En-
gineering, Indian Institute of Technology (ISM),
Dhanbad, Jharkhand, India
Archana S. Rao School of Basic and Applied Sci-
ences, Dayananda Sagar University, Bangalore,
India
xi
P.C. Rath ICAR-National Rice Research Insti-
tute, Cuttack, Odisha, India
Lopamudra Ray School of Biotechnology, Ka-
linga Institute of Industrial Technology (KIIT),
Bhubaneswar, Odisha, India; School of Law,
Kalinga Institute of Industrial Technology
(KIIT), Bhubaneswar, Odisha, India
Kiran K. Reddy ICAR-Directorate of Ground-
nut Research, Junagadh, Gujarat, India
Monire Mones Sardrodi Department of Plant
Breeding and Biotechnology, Faculty of Agri-
culture, University of Tabriz, Tabriz, Iran
Preeti Sengupta Amity Institute of Microbial
Technology, Amity University, Noida, Uttar
Pradesh, India
Efath Shahnaz Division of Plant Pathology,
Faculty of Horticulture, SKUAST-Kashmir, Sri-
nagar, J&K, India
Shilpi Sharma Department of Biochemical En-
gineering and Biotechnology, Indian Institute
of Technology Delhi, Hauz Khas, New Delhi,
India
Deepali Shukla Department of Biotechnology,
IPS Academy Indore, MP, India
Piyush Shukla Department of Rural Technolo-
gy, Guru Ghasidas Central University, Bilaspur,
CG, India
A.L. Singh ICAR-Directorate of Groundnut Re-
search, Junagadh, Gujarat, India
Poonam C. Singh Department of Plant microbe
interaction, CSIR-NBRI, Lucknow, UP, India
Prashant Kumar Singh Amity Institute of Mi-
crobial Technology, Amity University, Noida,
Uttar Pradesh, India
Bana Sravani Department of Plant Pathology
and Agricultural Microbiology, Post Graduate
Institute, Mahatma Phule Krishi Vidyapeet, Ra-
huri, Maharashtra, India
Ashmita Tandon Department of Plant Microbe
Interaction, CSIR-NBRI, Lucknow, UP, India
Stevenson Thabah Microbiology Laboratory,
Department of Biotechnology & Bioinformat-
ics, North-Eastern Hill University, Shillong,
Meghalaya, India
xii CONTRIBUTORS

Ajit Varma Amity Institute of Microbial
Technology, Amity University, Noida, Uttar
Pradesh, India
Ambarish S. Vidyarthi N.P.S Institute of Tech-
nology, GIC Campus, Pithoragarh, Uttara-
khand, India
M.K. Yadav Department of Plant Pathology,
Janta Collage, Bakewar, Etawah, UP, India
Manoj K. Yadav ICAR-National Rice Research
Institute, Cuttack, Odisha, India
 About the Editors
Professor Harikesh
Bahadur Singh is Distin-
guished ­Professor, Depart-
ment of Biotechnology,
GLA University, Mathura,
India. He has served as
Professor & Head, Depart-
ment of Mycology and
Plant Pathology at Institute of Agricultural
Sciences, Banaras Hindu University, India.
His major research focus is on bioinoculats,
biological control of plant pathogens and
nano-biotechnology. Prof. Singh has been
decorated with several prestigious National
awards and honors for his scientific contri-
butions in the field of organic farming and
translational agriculture, notable being CSIR
Technology Prize for Biological Sciences,
M.S. Swaminathan award by Society for
Plant Research, Vigyan Bharti Award, Prof.
V.P. Bhide Memorial Award, Society for Plant
Research, and many more. To his credit, he
has 20 U.S. patents which have been success-
fully transferred for commercial production
of biopesticides to several industrial houses
in India. He has been able to add more than
300 publications and 31 books Google Scholar
Citations- 10074; H-index- 52; i10- 182.
xiii
Dr. Anukool Vaishnav is
working as a Post-
Doctoral Scientist (Swiss
excellence fellow) at Uni-
versity of Zurich, Switzer-
land. He is also associated
with Department of Bio-
technology, GLA Univer-
sity, India as an Assistant
Professor. He has operated SERB-National
Postdoctoral Fellowship (NPDF) as Principal
Investigator at Department of Mycology and
Plant Pathology, Institute of Agricultural Sci-
ences, Banaras Hindu University, Varanasi,
India. He has >8 years of research experience
in plant-microbe interaction field. His research
is mainly focused on microbial mediated
plant protection against biotic and abiotic
stress, characterization of signaling molecules
and secondary metabolites (soluble and
VOCs) in plant holobionts. As an active
researcher, he has published more than
24 research and review articles along with
15 book chapters for reputed journals and
edited books. He has filed 9 Indian patents in
association with his research group. He is
editor of 2 Springer published books. He is an
editorial member of Current Genomics
Journal published by Bentham Science. He
has been awarded Swiss Excellence Post Doc-
toral Scholarship and several Young Scientist
Awards from different agencies.
 Preface
Zero hunger is a complicated task for scien-
tific community under this erratic climate
conditions, rapid population growth, and
with limited arable land. In the last two
years (2020 & 2021), the globe is encoun-
tered with the pandemic of Covid-19. This
outbreak causes declines and stagnation of
food trading and transportation between
nations that again affect global food supply.
To match with food demand, it is necessary
to apply such approaches that can increase
crop productivity and yield. The intensive
use of agrochemicals (fertilizers and pes-
ticides) has led to severe consequences,
including the deterioration of soil and the
environment. Increasing soil pollution and
adverse effects of agrochemicals on human
and animals built a pressure on agricul-
tural scientists to find alternate eco-friendly
approaches. In this context, application of
agriculturally important microorganisms
(AIMs) with the genera of Bacillus, Pseu-
domonas, Rhizobium, Burkholderia, Arhtobac-
ter, Trichoderma and Mycorrhiza spp. for the
purpose of biofertilizer and biopesticides is
a sustainable and eco-friendly alternative
of agrochemicals. The most consensual and
recent definition of these microbes states
“Plant Biostimulants”, which can be directly
applied to plants, seeds, and soil for sustain-
able agriculture production. These biostimu-
lants are part of a wide unknown microbial
diversity, constituting the rhizospheric, epi-
phytic, or endophytic microbiota, which are
xv
on the way to being cultured and preserved
at in-vitro conditions. This is an emerging
area in agrofarming, where a lot of evidences
demonstrate the potential of several AIMs in
promotion of plant growth and productivity
and also in disease management. However,
the mechanisms behind their efficacy and
interaction with soil biota and host plant
are still matter of discussion. The beneficial
microbes improve soil fertility and promote
plant growth through direct and indirect
processes. In direct process, these microbes
produce plant growth promoting hormones
and solubilize inorganic nutrients into the
soil that increase nutrient uptake efficiency
of plants leading to better growth and pro-
ductivity. Under indirect process, AIMs
work as biocontrol agent for different pests
and reduce pathogenicity in plants. There-
fore, these beneficial microbes can be used
as biofertilizers and biopesticides for cleaner
and healthier agricultural practices.
This book has reviewed the recent studies
on different agricultural important microbes
and emphasized on their future research
and application in agriculture. The chap-
ters contributed by prominent scientists will
be a ready source of information for young
researchers, industrialist and progressive
farmers.
Harikesh Bahadur Singh, Mathura, India
Anukool Vaishnav, Zürich, Switzerland
 C H A P T E R

1
Role of microorganism as new generation
plant bio-stimulants: An assessment
Deepali Shuklaa, Piyush Shuklab, Ashmita Tandonc, Poonam C.
Singhc, Jayandra Kumar Johric
aDepartment
of Biotechnology, IPS Academy Indore, MP, India, bDepartment of Rural
Technology, Guru Ghasidas Central University, Bilaspur, CG, India, cDepartment of Plant
Microbe Interaction, CSIR-NBRI, Lucknow, UP, India

1.1 Background

World’s largest population are dependent on plants or crop for their food requirements. For
the sustainable development of population, it is very essential that production of plants should
be raised more and more every time. In this context “Green revolution” played a vital role for
the production of rice, corn and wheat crop varieties. Moreover, different world governments
and organizations makes indispensable efforts to work on multiplication on the production
of crop to ensure the food provision (Abbott et al., 2018; Ortíz-Castro et al., 2012; Roberts and
Tasistro, 2012).
Studies suggest that for the increased productivity of crop, the role of nutrients like nitro-
gen, phosphorus, minerals and vitamins along with the plant health is very important. To
fulfilling demand of enhance crop productivity, in past decades chemical fertilizers market
were continuously growing globally (Singh et al., 2016). Intensive application of phosphorus
and nitrogen base chemical leads to a massive depletion of soil and water resources (Den
Herder et al., 2010), and it also cause global crisis of pollution. Enormous use of chemicals
is very harmful to human and environment too (Carvalho, 2017).
Other than use of chemical fertilizers for better crop yield climate change, emerging plant
pathogens, drought and stress is also a major challenge that affect agroecosystem (Rosenz-
weig and Tubiello, 2007). For surpass crop production, it is necessary to combat for all these
environmental hazards caused by the use of extensive chemical fertilizers and other climate,
plant disease and stress related problems for this some protocol and rules were established
for agropractices worldwide (Gomiero et al., 2011). Bio-stimulants gain popularity as addi-
tives to fertilizers which increase the efficacy of fertilizers by stimulating physiology of plant

New and Future Developments in Microbial Biotechnology and Bioengineering.

DOI: https://doi.org/10.1016/B978-0-323-85163-3.00016-8 1  Copyright © 2022 Elsevier Inc. All rights reserved.
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regarded as comprising two distinct types of nervous condition,
according as the loop formed by the two visceral nerve-cords is
twisted over itself, forming a figure of 8, or continues straight and
uncrossed. In the former case, we get the condition known as
streptoneurous, in the latter that as euthyneurous.[315] The
Euthyneura include the whole of the Opisthobranchiata[316] and
Pulmonata, the Streptoneura all the Prosobranchiata.

Fig. 101.—Nervous system of the Amphineura: A,

Proneomenia; B, Neomenia; C,
Chaetoderma; D, Chiton; c, cerebral ganglia;
l, l, lateral cords; pc, posterior commissure; s,
sublingual commissure or ring, with ganglia; v,
v, pedal cords. (Alter Hubrecht.)
The simplest form of nervous system in the euthyneurous
Gasteropoda occurs in the Opisthobranchiata. The cerebral, pleural,
and pedal ganglia tend to become concentrated in a ring-like form,
united by short commissures at the posterior end of the pharynx. The
visceral loop is in some cases long, and the two or three visceral
ganglia are then situated at its posterior extremity. The nervous
system of the Pulmonata is of a similar type, the visceral loop being
often much shorter, and tending to draw in towards the central group
of ganglia. The tentacular and optic nerves are, as usual, derived
from the cerebral ganglion, with which also the octocysts are
probably connected by rather long nerves. A pair of buccal ganglia
innervate the buccal mass, and are united by commissures with the
right and left cerebral ganglia. The osphradial nerve springs from
one of the ganglia on the visceral loop, the osphradium itself being
situated (in Limnaea) immediately above the pulmonary orifice and
adjacent to the anus (Fig. 102). This massing of the ganglia is still
better illustrated by the accompanying figure of Physa (Fig. 103), in
which the animal is represented as if transparent, so that the ganglia
and nerves are seen through the tissues.
Fig. 102.—I. Nervous system of Limnaea
stagnalis L. The oesophagus has been
cut and pulled forwards through the
nerve-collar, so as to expose the lower
surface of the buccal mass(dissected by
F. B. Stead)
B.M, buccal mass.
B.G, buccal, C.G, cerebral, Os.G,
osphradial, Pe.G, pedal ganglia.
Pl.G, pleural ganglia.
Op.N, optic, Os.N, osphradial, Te.N,
tentacular nerve.
Ot, otocyst; V.L, visceral loop.
R, rectum, dotted in to show its position
relative to the osphiadium.

II. Right side of the head of Limnaea

stagnalis. The overhanging flap of the
 mantle has been cut in the middle line,
and the right half twisted back, so as to
expose the pulmonary orifice, etc. The
points A A on the mantle edge were
continuous before the mantle was cut;
the line BA is part of the free edge of the
mantle.
An, anus; F, female generative orifice; J,
portion of jaw; M, male generative orifice
under right tentacle; Os, osphradium; P.O,
pulmonary orifice.
Of the streptoneurous Gasteropoda, the nervous system of
Fissurella and Haliotis shows distinct points of similarity to that of the
Amphineura. The pedal nerves are united by transverse
commissures throughout their entire length, while a double
commissure unites the cerebral ganglia to the mass from which the
pedal nerves proceed. In the great majority of the Streptoneura the
ganglia (except the visceral) are more concentrated and the
commissures are consequently much shorter. The accompanying
figure of Cyclostoma, in which the animal is represented as in that of
Physa just described, illustrates this grouping of the ganglia, the twist
of the visceral loop, and the position of the visceral ganglia at its
posterior end.
Fig. 103.—Nervous system of
Physa acuta Drap., showing
the massing of the ganglia at
the hinder end of the
pharynx: e, e, eyes; m,
mouth; m.l, m.l, mantle
lappets; o.f, female
generative orifice; o.m, male
generative orifice; os,
osphradium. (After Lacaze-
Duthiers.)
 Fig. 104.—Example of a
streptoneurous Gasteropod
(Cyclostoma elegans Drap.): c.g,
c.g, cerebral ganglia; e, e, eyes;
os, osphradium; ot, ot, otocysts;
p.g, p.g, pedal ganglia; pl.g, pl.g,
pleural ganglia; sp.g, supra-
intestinal ganglion; sb.g, sub-
intestinal ganglion; t.n, tentacle
nerve; v.g, visceral ganglion. (After
Lacaze-Duthiers.)
Scaphopoda.—In the Scaphopoda the nervous system
resembles that of the Pelecypoda. The cerebral and pleural ganglia
lie close together, while the pedal ganglia are placed in the anterior
part of the foot, connected with the cerebral ganglia by long
commissures; the visceral loop is rather long, and the two visceral
ganglia are adjacent to the anus.
Pelecypoda.—The nervous system in the Pelecypoda is the
simplest type in which well-marked ganglionic centres occur. The
ganglia are few, symmetrically placed, and are usually at a
considerable distance apart. There are, as a rule, three distinct pairs
of ganglia, the cerebral (cerebro-pleural), pedal, and visceral. The
cerebral are formed by the fusion of the cerebral and pleural ganglia,
which, however, in some cases (Protobranchiata) continue distinct.
[317] They lie above or on each side of the mouth, united by a
commissure of varying length. Another pair of commissures unites
them with the pedal ganglia, which are placed at the base of the foot,
and are usually very close together, sometimes (as in Anodonta)
becoming partially fused. The length of these commissures depends
upon the distance between mouth and foot; thus they are very long
in Mya and Modiola, and very short in Pecten. In cases where the
foot is rudimentary or becomes aborted through disuse (e.g. Ostrea),
the pedal ganglia may dwindle or disappear altogether. The visceral
ganglia are on the ventral side of the posterior adductor muscle,
beneath the rectum, and innervate the branchiae, osphradia, and the
whole of the visceral sac. A pair of cerebro-visceral commissures
traverses the base of the foot, surrounding it with a comparatively
short loop (compare Fig. 106, c.v.c´), while a long commissure,
which runs round the entire edge of the mantle, and supplies
branching nerves to the mantle border and siphons (Fig. 106, c.v.c),
may also connect the visceral and cerebral ganglia.
 Fig. 105.—Nervous system of
Pelecypoda: A, Teredo; B,
Anodonta; C, Pecten; a, a,
cerebral ganglia; b, pedal
ganglia; c, visceral
ganglia. (After
Gegenbaur.)
Cephalopoda.—In the Cephalopoda the concentration of ganglia
attains its maximum, and may perhaps be regarded as approaching
the point at which a definite brain may be said to exist. Another point
of distinction is the formation of special small ganglia upon the
nerve-cords in different parts of the body. In the Tetrabranchiata
(Nautilus) the cerebral and pedal ganglia form a broad ring which
surrounds the oesophagus, the former giving out the optic nerves,
with their special optic ganglion, and a pair each of buccal and
pharyngeal ganglia, the latter the nerves of the arms and funnel. The
visceral loop is still present in the form of a separate band, which
innervates the branchiae, osphradia, and viscera generally, forming
a special genital ganglion in connexion with the reproductive organs.
The principal ganglia of the Dibranchiata are still more concentrated,
even the visceral loop being possibly united with the rest in forming
an unbroken mass in which scarcely any trace of commissures can
be detected. The pedal ganglion becomes separated into two
portions, one of which innervates the arms, the other the funnel. Two
peculiar ganglia (the stellate ganglia) supply a number of branching
nerves to the mantle.

Fig. 106.—Nervous system of Cardium edule L.: a.m, anterior adductor

muscle; br, branchiae; br.n, branchial nerve; c.g, c.g, cerebral ganglia;
c.p.c, cerebro-pedal commissure; c.v.c’, cerebro-visceral commissure;
c.v.c, cerebro-visceral commissure of mantle; l.p, labial palps: m, mouth;
p.g, pedal ganglion; p.m, posterior adductor muscle; v.g, visceral ganglion.
(After Drost, × 3.)

E. L. Bouvier, Système nerveux, morphologie générale et

classification des Gastéropodes prosobranches: Ann. Sc. Nat. Zool.
(7), iii. 1887, pp. 1–510.
J. Brock, Zur Neurologie der Prosobranchier: Zeit. wiss. Zool. xlviii.
1889, pp. 67–83.
 O. Bütschli, Bemerkungen über die wahrscheinliche Herleitung der
Asymmetrie der Gasteropoda, etc.: Morph. Jahrb. xii. 1886, pp. 202–
222.
B. Haller, Zur Kenntniss der Muriciden. I. Anatomie des
Nervensystems: Denksch. Math. Nat. Kl. Ak. Wien, xlv. 1882, pp. 87–
106.
„ Untersuchungen über marine Rhipidoglossen. II. Textur des„
Centralnervensystems und seiner Hüllen: Morph. Jahrb. xi. 1885, pp.
319–436.
H. Grenadier, Abhandlungen zur vergleichenden Anatomie des
Auges: Abh. Naturf. Gesell. Halle, xvi. 1884, pp. 207–256; xvii. 1886,
pp. 1–64.
A. P. Henchman, The Origin and Development of the Central
Nervous System in Limax maximus: Bull. Mus. C. Z. Harv. xx. 1890,
pp. 169–208.
V. Hensen, Ueber das Auge einiger Cephalophoren: Zeit. wiss.
Zool. xv. 1865, pp. 157–242.
C. Hilger, Beiträge zur Kenntniss des Gasteropodenauges: Morph.
Jahrb. x. 1885, pp. 352–371.
Lacaze-Duthiers, Otocystes ou Capsules auditives des
Mollusques (Gastéropodes): Arch. Zool. Exp. Gén. i. 1872, pp. 97–
166.
„ „ Du système nerveux des Mollusques gastéropodes
pulmonés aquatiques: ibid. pp. 437–500.
P. Pelseneer, Recherches sur le système nerveux des Ptéropodes:
Arch. Biol. vii. 1887, pp. 93–130.
„ Sur la valeur morphologique des bras et la composition du
système nerveux central des Cephalopodes: Arch. Biol. viii. 1888, pp.
723–756.
H. Simroth, Ueber die Sinneswerkzeuge unserer einheimische
Weichthiere: Zeit. wiss. Zool. xxvi. 1876, pp. 227–348.
J. W. Spengel, Die Geruchsorgane und das Nervensystem der
Mollusken: Zeit. wiss. Zool. xxxv. 1881, pp. 333–383.
 CHAPTER VIII
THE DIGESTIVE ORGANS, JAW, AND RADULA: EXCRETORY ORGANS

The digestive tract, or, as it is often termed, the alimentary canal

or gut, is a very important feature of the Mollusca. It may be
regarded as consisting of the following parts: (1) a mouth or oral
aperture: (2) a throat or pharynx; (3) an oesophagus, leading into (4)
a stomach, (5) an intestine and rectum, ending in (6) an anus.
The primitive positions of mouth and anus were presumably at the
anterior and posterior ends of the animal, as in the Amphineura and
symmetrical Mollusca generally. But the modifications of original
molluscan symmetry, which have already been referred to (p. 154,
compare pp. 245, 246), have resulted in the anus becoming, in the
great majority of Gasteropoda, twisted forward, and occupying a
position on some point in the right side in dextral, and in the left in
sinistral species.
The process of digestion, as the food passes from one end of the
tract to the other, is performed by the aid of the secretions of various
glands, which open into the alimentary canal at different points in its
course. The principal of these are the salivary glands, situated on the
pharynx and oesophagus, and the liver, biliary or hepatic gland,
connecting with the stomach. With these may be considered the anal
and ink-glands, which, in certain genera, connect with the terminal
portion of the rectum.
1. The mouth is generally, as in the common snail and periwinkle,
placed on the lower part of the head, and may be either a mere
aperture, circular or semicircular, in the head-mass, or, as is more
usual, may be carried on a blunt snout (compare Fig. 6, p. 10, and
Fig. 68, p. 159), which is capable of varying degrees of protrusion.
From the retractile snout has doubtless been derived the long
proboscis which is so prominent a feature of many genera (compare
Figs. 1, B, and 99), and in some (e.g. Mitra, Dolium) attains a length
exceeding that of the whole body. As a rule, Mollusca provided with
a proboscis are carnivorous, while those whose mouth is on the
surface of the head are Vegetable feeders, but this rule is by no
means invariable. The mouth is thickened round the aperture into
‘lips,’ which are often extensile, and appear capable of closing upon
and grasping the food. In the Pelecypoda the mouth is furnished, on
each side, with a pair of special external lobes, the ‘labial palps,’
which appear to be of a highly sensitive nature, and whose object it
is to collect, and possibly to taste, the food before it passes into the
mouth.
2. The Pharynx, Jaws, and Radula.—Immediately behind the lips
the mouth opens into the muscular throat, pharynx, or buccal mass.
The pharynx of the Glossophora, i.e. of the Gasteropoda,
Scaphopoda, and Cephalopoda, is distinguished from that of the
Pelecypoda,[318] by the possession of two very characteristic organs
for the rasping or trituration of food before it reaches the oesophagus
and stomach. These are (a) the jaw or jaws, and (b) the radula,[319]
odontophore, or lingual ribbon. The jaws bite the food, the radula
tears it up small before it passes into the stomach to undergo
digestion. The jaws are not set with teeth like our own; roughly
speaking, the best idea of the relations of the molluscan jaw and
radula may be obtained by imagining our own teeth removed from
our jaws and set in parallel rows along a greatly prolonged tongue.
[320]

In nearly all land Pulmonata the jaw is single, and is placed

behind the upper lip. If a common Helix aspersa be observed
crawling up the inside of a glass jar, or feeding on some succulent
leaf, the position and action of the jaw can be readily discerned. It
shows very black when the creature opens its mouth, and under its
operation the edge of a lettuce leaf shows a regular series of little
curved indentations, in shape not unlike the semicircular bites
inflicted by a schoolboy upon his bread and butter. The jaw of Helix
(Fig. 107, B) is arched in shape, and is strengthened by a number of
projecting vertical ribs. That of Limax (A) is straighter, and is slightly
striated, without vertical ribs. In Bulimulus (C) the arch of the jaw is
very conspicuous, and the upper edges are always denticulated; in
Orthalicus there is a central triangular plate with a number of
overlapping plates on either side; in Succinea (E) there is a large
square accessory plate above the jaw proper. The form of the jaw is
peculiar not only to the genus but to the species as well. Thus the
jaw of H. aspersa is specifically distinct from that of H. pomatia, and
that of H. nemoralis is distinct from both. Wiegmann has
observed[321] that in young Arion, Limax, and Helix, the jaw consists
of two pieces, which coalesce by fusion in the adult, thus indicating a
stage of development in advance of the double jaw which is found in
most of the non-pulmonate Mollusca. In all fresh-water Pulmonata
there are two small accessory side plates besides the jaw proper
(Fig. 107, F).

Fig. 107.—Jaws of various Pulmonata: A, Limax

(gagates Drap., Lancashire, × 15); B, Helix
(acutissima Lam., Jamaica, × 15); C,
Bulimulus (depictus Reeve, Venezuela, × 20);
D, Achatina (fulica Fér., Mauritius, × 7); E,
Succinea (elegans Riss., Aral District, × 30);
F, Limnaea (stagnalis L., Cambridge, × 30).
Nearly all the non-carnivorous Prosobranchiata, land, fresh-water,
and marine alike, are provided with two large lateral jaws. Many of
these are sculptured with the most elaborate patterns, and appear to
be furnished with raised teeth, like a file. In the Nudibranchiata the
jaws are of great size and beauty of ornamentation (Fig. 109).

Fig. 108.—Jaws of A, Triton australis Lam., Sydney; B, Ampullaria

fasciata Reeve, Demerara; C, Calliostoma punctulatum Mart.,
New Zealand; D, Cyclophorus atramentarius Sowb., Sanghir; all
× 15.
 Fig. 109.—Jaws of A, Chromodoris gracilis Iher., × 15; B,
Scyllaea pelagica L., × 7; C, Pleurobranchus plumula
Mont., × 10; D, Pleurobranchaea Meckelii Lam., × 5/2.

The carnivorous genera, whether marine (e.g. Conus, Murex,

Buccinum, Nassa) or land (e.g. Testacella, Glandina, Streptaxis,
Ennea), are entirely destitute of jaws, the reason probably being that
in all these cases the teeth of the radula are sufficiently powerful to
do the work of tearing up the food without the aid of a masticatory
organ as well. Jaws are also wanting in the Heteropoda, and in many
of the Nudibranchiata and Tectibranchiata.
In the Cephalopoda the jaws, or ‘beaks,’ as they are called, are
most formidable weapons of attack. In shape they closely resemble
the beaks of a parrot, but the hook on the dorsal side of the mouth
does not, as in birds, close over the lower hook, but fits under it.
Powerful muscles govern these mandibles, which must operate with
immense effect upon their prey (Fig. 110).
Fig. 110.—Jaws of Sepia:
A, in situ within the
buccal mass, several
of the arms having
been cut away; B,
removed from the
mouth and slightly
enlarged.
 Fig. 111.—Patella vulgata L.,
showing the normal position
of the radula, which is
doubled back in a bow; the
shell has been removed, and
the whole visceral mass is
turned forward, exposing the
dorsal surface of the
muscular foot: gr,
longitudinal groove on this
surface; i, i, intestine; l, liver;
m, m, mantle edge; mu,
muscles (cut through)
fastening the visceral mass
to the upper sides of the foot;
ov, ovary; r, radula; u.f,
upper or dorsal surface of
the foot.

The Radula.[322]—When the food has passed beyond the

operation of the jaw, it comes within the province of the radula, the
front part of which perhaps co-operates to a certain extent with the
jaw in performing the biting process. The function of the radula as a
whole is to tear or scratch, not to bite; the food passes over it and is
carded small, the effect being very much the same as if, instead of
dragging a harrow over the surface of a field, we were to turn the
harrow points upwards, and then drag the field over the harrow.
The radula itself is a band or ribbon of varying length and breadth,
formed of chitin, generally almost transparent, sometimes beautifully
coloured, especially at the front end, with red or yellow.[323] It lies
enveloped in a kind of membrane, in the floor of the mouth and
throat, being quite flat in the forward part, but usually curving up so
as to line the sides of the throat farther back, and in some cases
eventually forming almost a tube. The upper surface, i.e. the surface
over which the food passes, is covered with teeth of the most varied
shape, size, number, and disposition, which are almost invariably
arranged in symmetrical rows. These teeth are attached to the
cartilage on which they work by muscles which serve to erect or
depress them; probably also the radula as a whole can be given a
forward or backward motion, so as to rasp or card the substances
which pass over it.
The teeth on the front part of the radula are often much worn (Fig.
112), and probably fall away by degrees, their place being taken by
others successively pushed up from behind. At the extreme hinder
end of the radula the teeth are in a nascent condition, and there are
often as many as a dozen or more scarcely developed rows. Here,
too, lie the cells from which the teeth are originally formed.
The length and breadth of the radula vary greatly in different
genera. In Littorina it is very narrow, and several times the length of
the whole animal. It is kept coiled away like a watch-spring at the
back of the throat, only a small proportion of the whole being in use. I
have counted as many as 480 rows in the common Littorina littorea.
In Patella it is often longer than the shell itself, and if the radula of a
large specimen be freshly extracted and drawn across the hand, the
action of the hooks can be plainly felt. In Aerope, the Turbinidae
generally, and Haliotis it is very large. In Turritella, Aporrhais,
Cylichna, Struthiolaria, and the Cephalopoda it is small in proportion
to the size of the animal. In the Pulmonata generally it is very broad,
the length not exceeding, as a rule, thrice the breadth; in most other
groups the breadth is inconsiderable, as compared to the length.
 The Radula is wanting in two families of Prosobranchiata, the
Eulimidae and Pyramidellidae, which are consequently grouped
together as the section Gymnoglossa. It is probable that in these
cases the radula has aborted through disuse, the animals having
taken to a food which does not require trituration. Thus several
genera contained in both these families are known to live
parasitically upon various animals—Holothurians, Echinoderms, etc.
—nourishing themselves on the juices of their host. In some cases,
the development of a special suctorial proboscis compensates for
the loss of radula (see pp. 76–77). In Harpa there is no radula in the
adult, though it is present in the young form. No explanation of this
fact has yet been given. It is also absent in the Coralliophilidae, a
family closely akin to Purpura, but invariably parasitic on corals, and
probably nourished by their exudations. There is no radula in
Entoconcha, an obscure form parasitic on the blood-vessels of
Synapta, or in Neomenia, a genus of very low organisation, or in the
Tethyidae, or sea-hares, or in one or two other genera of
Nudibranchiata.
 Fig. 112.—Example of a front
portion of a radula
(Cantharus ringens Reeve,
Panama), much worn by use.
× 70.
The number of teeth in the radula varies greatly. When the teeth
are very large, they are usually few in number, when small, they are
very numerous. In the carnivorous forms, as a rule, the teeth are
comparatively few and powerful, while in the phytophagous genera
they are many and small. Large hooked and sickle-shaped teeth,
sometimes furnished with barbs like an arrow-head, and poison-
glands, are characteristic of genera which feed on flesh; vegetable
feeders, on the contrary, have the teeth rounded, and blunter at the
apex, or, if long and narrow, so slender as to be of comparatively
little effect. Genera which are normally vegetarian, but which will,
upon occasion, eat flesh, e.g. Limax and Hyalinia, exhibit a form of
teeth intermediate between these two extremes (see Fig. 140, A).
In Chaetoderma there is but one tooth. In Aeolis coronata there
are about 17, in A. papillosa and Elysia viridis about 19, in Glaucus
atlanticus about 21, in Fiona nobilis about 28. In the common whelk
(Buccinum undatum) there are from 220 to 250, in the common
periwinkle about 3500. As many as 8343 have been counted in