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Plant Transcription Factors
Contribution in Development,
Metabolism, and Environmental Stress
This page intentionally left blank
Plant Transcription Factors
Contribution in Development,
Metabolism, and Environmental Stress

Edited by
Vikas Srivastava
Department of Botany, School of Life Sciences, Central University of Jammu,
Samba, Jammu and Kashmir (UT), India

Sonal Mishra
Department of Botany, School of Life Sciences, Central University of Jammu,
Samba, Jammu and Kashmir (UT), India

Shakti Mehrotra
Department of Biotechnology, Institute of Engineering and Technology,
Lucknow, Uttar Pradesh, India

Santosh Kumar Upadhyay


Department of Botany, Panjab University, Chandigarh (UT), India
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Acquisitions Editor: Nancy J. Maragioglio
Editorial Project Manager: Lena Sparks
Production Project Manager: Omer Mukthar
Cover Designer: Miles Hitchen

Typeset by MPS Limited, Chennai, India


Contents

List of contributors ............................................................................................................................ xvii


About the editors .............................................................................................................................. xxiii
Preface ............................................................................................................................................... xxv

Section I Plant transcription factors (TFs) and general aspects


CHAPTER 1 Plant transcription factors: an overview of their role in
plant life ................................................................................................ 3
Aksar Ali Chowdhary, Sonal Mishra, Shakti Mehrotra,
Santosh Kumar Upadhyay, Diksha Bagal and Vikas Srivastava
1.1 Introduction ................................................................................................................ 3
1.2 Transcription factors and plant life............................................................................ 4
1.3 Transcription factors and stress responses................................................................. 6
1.4 Transcription factors and secondary metabolism ...................................................... 9
1.5 Conclusion ................................................................................................................ 10
References................................................................................................................. 11

CHAPTER 2 Adaptation of millets to arid land: a special perspective


of transcription factors ....................................................................... 21
Alka Bishnoi, Pooja Jangir and Praveen Soni
Highlights.................................................................................................................. 21
Abbreviations............................................................................................................ 21
2.1 Introduction .............................................................................................................. 22
2.2 Distribution of arid land in India and world ........................................................... 23
2.3 Millets: climate-smart nutri-cereals ......................................................................... 24
2.4 Stress as a limiting factor for crops in the arid zones............................................. 27
2.5 Responses of millets to abiotic stresses................................................................... 28
2.6 Transcription factors: smart regulators of stress tolerance in millets ..................... 29
2.6.1 WRKY.......................................................................................................... 30
2.6.2 DOF .............................................................................................................. 31
2.6.3 ERF/DREB................................................................................................... 40
2.6.4 NAC ............................................................................................................. 41
2.6.5 bHLH............................................................................................................ 41
2.6.6 ASR .............................................................................................................. 42
2.6.7 bZIP.............................................................................................................. 42
2.6.8 MYB............................................................................................................. 42
2.6.9 SBPs ............................................................................................................. 43

v
vi Contents

2.6.10 Other transcription factors ........................................................................... 43


2.7 Harnessing the potential of millet transcription factors .......................................... 43
2.8 Conclusion and future perspectives ......................................................................... 46
Acknowledgments .................................................................................................... 47
Declaration of competing interests .......................................................................... 47
Author contribution .................................................................................................. 47
References................................................................................................................. 47

Section II Plant TFs and development .................................................. 61


CHAPTER 3 Plant transcription factors and root development............................. 63
Rekha Chouhan, Abhilek Kumar Nautiyal,
Nancy Sharma and Sumit G. Gandhi
3.1 Introduction .............................................................................................................. 63
3.2 Plant root architecture and development ................................................................. 64
3.3 Transcription factors involved in plant root development ...................................... 64
3.3.1 Root apical meristem ..................................................................................... 64
3.3.2 Lateral roots ................................................................................................... 67
3.3.3 Root hair......................................................................................................... 69
3.4 Conclusion ................................................................................................................ 71
Acknowledgments .................................................................................................... 72
References................................................................................................................. 72

CHAPTER 4 The roles of transcription factors in the development of


plant meristems .................................................................................. 77
Qingkun Dong and Cui Zhang
4.1 Introduction .............................................................................................................. 77
4.2 Shoot apical meristem.............................................................................................. 77
4.3 Axillary meristem..................................................................................................... 79
4.4 Flower meristem....................................................................................................... 82
4.5 Intercalary meristem................................................................................................. 84
4.6 Conclusion and future perspectives ......................................................................... 84
Acknowledgments .................................................................................................... 88
Author contributions................................................................................................. 88
References................................................................................................................. 88

CHAPTER 5 Transcription factors and their role in leaf senescence .................. 93


Jeremy Dkhar and Asosii Paul
5.1 Introduction .............................................................................................................. 93
Contents vii

5.2 Identification of transcription factor families in senescing leaf


transcriptome ............................................................................................................ 94
5.3 Characterization of leaf senescence related TFs families ....................................... 98
5.3.1 No apical meristem (NAM), ATAF1/2, CUP-shaped
cotyledon 2 (CUC2) (NAC) TF................................................................... 98
5.3.2 WRKY TF.................................................................................................. 104
5.3.3 APETALA2/Ethylene-responsive element binding protein
(AP2/EREBP) superfamily ........................................................................ 107
5.3.4 Basic helix-loop-helix (bHLH) TFs .......................................................... 108
5.3.5 MYB TFs ................................................................................................... 110
5.3.6 Auxin response factor and Auxin/INDOLE-3-acetic acid TFs................. 110
5.3.7 DNA binding-with-one-finger (DOF) proteins ......................................... 112
5.3.8 PSEUDO-response regulators TF .............................................................. 113
5.3.9 VQ TF family............................................................................................. 113
5.3.10 Basic leucine zipper (bZIP) TFs................................................................ 113
5.3.11 Homodomain-leucine zipper (HD-ZIP) TFs ............................................. 114
5.3.12 Plant A/T-rich sequence and zinc-binding protein (PLATZ) TF ............. 115
5.3.13 Growth-regulating factors (GRFS) and GRF-interacting
factors (GIFS) ............................................................................................ 115
5.3.14 Teosinte branched 1, Cycloidea, and proliferating cell nuclear
antigen binding factor (TCP) TFS............................................................. 116
5.3.15 Homeobox (HB) TFs ................................................................................. 116
5.3.16 C3H (Zn) TFs............................................................................................. 117
5.3.17 GRAS TFs.................................................................................................. 118
5.3.18 CCAAT box-binding TFs .......................................................................... 119
5.3.19 Heat shock factor TFs................................................................................ 119
5.3.20 MADS TFs ................................................................................................. 119
5.3.21 GOLDEN 2, ARR-B, PSR 1 (GARP) family TFs.................................... 120
5.3.22 TRIHELIX TFs .......................................................................................... 120
5.3.23 Arabidopsis response regulator TFs .......................................................... 121
5.3.24 Lateral organ boundaries/asymmetric leaves 2 ......................................... 122
5.3.25 Early flowering 3 (ELF3) TF .................................................................... 122
5.3.26 Ethylene insensitive 3 (EIN3)-like (EIL) TFS .......................................... 122
5.3.27 Brinsensitive 1 (BRI1)-EMS-Suppressor1 (BES1) TF ............................. 123
5.3.28 Calmodulin-binding transcription activator............................................... 123
5.3.29 TIFY TFs.................................................................................................... 124
5.3.30 B-box zinc finger TFs................................................................................ 124
5.4 Conclusion .............................................................................................................. 125
Acknowledgments .................................................................................................. 125
References............................................................................................................... 126
viii Contents

CHAPTER 6 Plant transcription factors in light-regulated development


and UV-B protection.......................................................................... 139
Deeksha Singh, Nevedha Ravindran, Nikhil Job,
Puthan Valappil Rahul, Lavanya Bhagavatula and Sourav Datta
6.1 Introduction ............................................................................................................ 139
6.1.1 Transcription factors families involved in light-regulated processes ......... 140
6.1.2 Transcription factors associated with visible light-mediated
development in plants .................................................................................. 143
6.1.3 Transcriptional regulation of UV-B signaling in plants ............................. 145
6.1.4 Structural and functional evolution of light-responsive plant
transcription factors ..................................................................................... 146
6.1.5 Role of light-regulated transcription factors in other signaling
pathways....................................................................................................... 148
6.2 Conclusion .............................................................................................................. 149
References............................................................................................................... 149

CHAPTER 7 Tomato fruit development through the perspective of


transcription factors ......................................................................... 159
Vigyasa Singh, Dharitree Phukan and Ujjal Jyoti Phukan
7.1 Introduction ............................................................................................................ 159
7.2 Transcription factors in tomato.............................................................................. 160
7.3 MYB transcription factors ..................................................................................... 161
7.4 MADS transcription factor..................................................................................... 163
7.5 Other transcription factors ..................................................................................... 165
7.6 Conclusion and future perspectives ....................................................................... 166
Acknowledgment .................................................................................................... 167
Conflict of interest.................................................................................................. 167
References............................................................................................................... 167

CHAPTER 8 Plant transcription factors and nodule development ...................... 175


Jawahar Singh and Praveen Kumar Verma
8.1 Introduction ............................................................................................................ 175
8.2 CCaMK/CYCLOPS complex................................................................................. 177
8.3 AP2-ERF transcription factor (ERN1 and ERN2) ................................................ 179
8.4 GRAS transcription factor ..................................................................................... 180
8.4.1 Nodulation signaling pathway 1/2 (NSP1 and NSP2) ................................ 180
8.5 SymSCL1 ............................................................................................................... 181
8.6 NIN and NIN-like proteins .................................................................................... 181
8.7 Structure of NIN and NLPs ................................................................................... 181
Contents ix

8.8 Regulation of NIN for rhizobial infection in the epidermis by CYCLOPS ......... 182
8.9 Regulation of NIN by cytokinin-response elements for cell
divisions in the pericycle ....................................................................................... 182
8.10 NIN: a master regulator of nodulation .................................................................. 183
8.11 NIN as a negative regulator in systemic control of nodulation ............................ 183
8.12 NIN as a positive regulator in systemic control of nodulation ............................. 184
8.12.1 Lob-domain protein16 ............................................................................... 185
8.12.2 Nodulation pectate lyase 1......................................................................... 185
8.13 Rhizobium-directed polar growth .......................................................................... 186
8.14 Nuclear factor Y..................................................................................................... 186
8.14.1 Short internodes/stylish.............................................................................. 187
8.15 Conclusion and future perspectives ....................................................................... 188
Acknowledgments .................................................................................................. 189
Declaration of competing interest .......................................................................... 189
Contribution ............................................................................................................ 189
References............................................................................................................... 189

Section III Plant TFs and metabolism .................................................. 197


CHAPTER 9 The regulatory aspects of plant transcription factors in
alkaloids biosynthesis and pathway modulation............................. 199
Pravin Prakash, Rituraj Kumar and Vikrant Gupta
Abbreviations.......................................................................................................... 199
9.1 Introduction ............................................................................................................ 199
9.2 Plant transcription factor families involved in alkaloid biosynthesis
regulation................................................................................................................ 201
9.2.1 APETALA2/ethylene response factor ......................................................... 201
9.2.2 Basic helix-loop-helix .................................................................................. 205
9.2.3 Basic leucine zipper ..................................................................................... 206
9.2.4 Cys2/His2-type (transcription factor IIIA-type) zinc-finger
protein family/Zinc-finger Catharanthus protein (ZCT) family................. 207
9.2.5 Myeloblastosis.............................................................................................. 207
9.2.6 WRKY.......................................................................................................... 208
9.2.7 Other transcription factors ........................................................................... 209
9.3 Transcription factor-mediated modulation of alkaloid biosynthesis pathways ........... 209
9.3.1 Overexpression............................................................................................. 209
9.3.2 Downregulation............................................................................................ 210
9.3.3 CRISPR/Cas-mediated genome editing....................................................... 211
9.4 Conclusions ............................................................................................................ 212
References............................................................................................................... 212
x Contents

CHAPTER 10 Plant transcription factors and flavonoid metabolism .................... 219


Rekha Chouhan, Garima Rai and Sumit G. Gandhi
10.1 Introduction ............................................................................................................ 219
10.2 Plant flavonoids, major subclasses, and biosynthesis ........................................... 220
10.3 Transcription factor families associated with plant flavonoid metabolism .......... 221
10.3.1 Role of basic-helix-loop-helix transcription factors in plant
flavonoid metabolism................................................................................. 222
10.3.2 MYB transcription factor family and plant flavonoid metabolism .......... 224
10.3.3 WD40 transcription factors and plant flavonoid metabolism................... 225
10.3.4 Role of basic leucine-zipper transcription factors in plant
flavonoid metabolism................................................................................. 226
10.3.5 Role of WRKY transcription factors in plant flavonoid metabolism............ 227
10.4 Conclusions ............................................................................................................ 227
Acknowledgments .................................................................................................. 227
References............................................................................................................... 227

CHAPTER 11 Demystifying the role of transcription factors in plant


terpenoid biosynthesis...................................................................... 233
Ajay Kumar, Parul Sharma, Rakesh Srivastava and
Praveen Chandra Verma
11.1 Introduction ............................................................................................................ 233
11.2 Biosynthesis of terpenoids ..................................................................................... 234
11.2.1 Biosynthesis of basic terpenoids precursor (MVA and MEP pathway)........ 235
11.2.2 Biosynthesis of isoprenoid intermediates .................................................. 235
11.2.3 Biosynthesis of terpenes by terpene synthases.......................................... 237
11.3 Regulation of terpenoids ........................................................................................ 239
11.3.1 WRKY........................................................................................................ 241
11.3.2 MYB........................................................................................................... 241
11.3.3 bHLH (basic helixloophelix) ............................................................... 242
11.3.4 AP2/ERF .................................................................................................... 242
11.3.5 bZIP ............................................................................................................ 243
11.3.6 SPL, YABBY, and other TFs .................................................................... 244
11.4 Conclusion .............................................................................................................. 244
Acknowledgment .................................................................................................... 244
References............................................................................................................... 244

CHAPTER 12 The regulatory circuit of iron homeostasis in rice:


a tale of transcription factors .......................................................... 251
Pooja Kanwar Shekhawat, Hasthi Ram and Praveen Soni
Highlights................................................................................................................ 251
Contents xi

Abbreviations.......................................................................................................... 251
12.1 Introduction ............................................................................................................ 252
12.2 Iron uptake and transport ....................................................................................... 253
12.3 Major transcription factors involved in iron homeostasis..................................... 255
12.3.1 Regulation of Fe deficiency....................................................................... 255
12.3.4 Regulation of Fe toxicity ........................................................................... 258
12.4 Regulation of the regulators................................................................................... 258
12.4.1 Epigenetic regulation ................................................................................. 259
12.4.2 Regulation at the transcriptional level....................................................... 259
12.4.3 Regulation at the post-transcriptional level............................................... 259
12.4.4 Regulation at the post-translational level .................................................. 259
12.4.5 Regulation by plant hormones................................................................... 260
12.5 Conclusion and future perspectives ....................................................................... 260
Acknowledgments .................................................................................................. 262
Author contribution ................................................................................................ 262
References............................................................................................................... 262

Section IV Plant TFs and Stress........................................................... 269


CHAPTER 13 Impact of transcription factors in plant abiotic stress:
a recent advancement for crop improvement.................................. 271
Divya Chauhan, Devendra Singh, Himanshu Pandey, Zeba Khan,
Rakesh Srivastava, Vinay Kumar Dhiman and Vivek Kumar Dhiman
13.1 Introduction ............................................................................................................ 271
13.2 Regulatory function of transcription factors in response to abiotic stress............ 271
13.3 ABA signaling pathway ......................................................................................... 272
13.4 JA signaling pathway ............................................................................................. 274
13.5 Transcription factors involved in abiotic stress tolerance..................................... 275
13.5.1 MYB TFs ................................................................................................... 275
13.5.2 NAC TFs .................................................................................................... 275
13.5.3 AP2/ERF TFs ............................................................................................. 276
13.5.4 WRKY TFs ................................................................................................ 277
13.6 Conclusion .............................................................................................................. 280
References............................................................................................................... 280

CHAPTER 14 Plant transcription factors and temperature stress ........................ 287


Tingting Zhang and Yang Zhou
14.1 Effect of temperature stress on plant growth ........................................................ 287
14.1.1 Effect of high-temperature stress on plants............................................... 287
xii Contents

14.1.2 Effect of low-temperature stress to plants................................................. 288


14.2 Transcription factors involved in response to temperature stress......................... 289
14.2.1 HSF transcription factor............................................................................. 289
14.2.2 MYB transcription factor........................................................................... 290
14.2.3 AP2/ERF transcription factors................................................................... 291
14.2.4 WRKY transcription factors ...................................................................... 293
14.3 Conclusions and perspectives ................................................................................ 294
Acknowledgments .................................................................................................. 294
References............................................................................................................... 294

CHAPTER 15 Plant transcription factors and osmotic stress ............................... 301


Tingting Zhang and Yang Zhou
15.1 Effects of osmotic stress on plants and its regulatory mechanism ....................... 301
15.1.1 Stomatal closure......................................................................................... 301
15.1.2 Osmotic regulation mechanism ................................................................. 302
15.1.3 Mechanism of ROS generation and scavenging ....................................... 302
15.1.4 ABA signaling pathway............................................................................. 303
15.2 Transcription factors are involved in regulating osmotic stress ........................... 304
15.2.1 Osmotic stress caused by salt stress .......................................................... 304
15.2.2 Osmotic stress caused by drought stress ................................................... 305
15.2.3 Osmotic stress caused by low temperature ............................................... 305
15.3 Conclusions and perspectives ................................................................................ 306
Acknowledgments .................................................................................................. 307
References............................................................................................................... 307

CHAPTER 16 Transcriptional regulation of drought stress stimulus:


challenges and potential for crop improvement ............................. 313
Gyanendra K. Rai, Gayatri Jamwal, Isha Magotra,
Garima Rai and R.K. Salgotra
16.1 Introduction ............................................................................................................ 313
16.2 Regulatory role of transcription factors in dry spell tolerance ............................. 314
16.3 Transcription factor and their mechanisms under drought stress ......................... 316
16.3.1 DNA binding with one finger (DOF) ........................................................ 316
16.3.2 WRKY transcription factor........................................................................ 318
16.3.3 Heat shock factor ....................................................................................... 319
16.3.4 Nuclear Factor (NF-Ys) ............................................................................. 320
16.3.5 TCP transcription factor family................................................................. 320
16.3.6 AP2/ERBP.................................................................................................. 322
16.3.7 AREB/ABF family..................................................................................... 323
Contents xiii

16.3.8 NAC transcription factors.......................................................................... 324


16.3.9 MYB/MYC transcription factors ............................................................... 325
16.4 Conclusion and future prospects............................................................................ 327
References............................................................................................................... 327

CHAPTER 17 Plant response to heavy metal stress: an insight into the


molecular mechanism of transcriptional regulation ....................... 337
Mehali Mitra, Puja Agarwal and Sujit Roy
17.1 Introduction ............................................................................................................ 337
17.2 Toxic effects of heavy metals in plants................................................................. 340
17.3 Plant signaling in response to heavy metal stress ................................................. 341
17.4 MAPK signaling under heavy metal stress ........................................................... 343
17.5 Calciumcalmodulin signaling pathway under heavy metal stress ..................... 344
17.6 Hormone signaling in response to heavy metal stress .......................................... 345
17.7 Reactive oxygen species production and its role in heavy metal stress ............... 346
17.8 Role of transcription factors in heavy metal resistance regulation....................... 347
17.9 The MYB-family transcription factors under HM stress ...................................... 348
17.10 The WRKY-family transcription factors under HM stress ................................... 352
17.11 The bZIP-family transcription factors under HM stress ....................................... 353
17.12 The AP2/ERF/DREB-family transcription factors under HM stress .................... 354
17.13 Conclusion and future perspectives ....................................................................... 355
Acknowledgments .................................................................................................. 355
References............................................................................................................... 355

CHAPTER 18 Plant transcription factors and salt stress ...................................... 369


Tingting Zhang and Yang Zhou
18.1 Effects of salt stress on plants ............................................................................... 369
18.1.1 Osmotic stress ............................................................................................ 369
18.1.2 Ion stress .................................................................................................... 369
18.1.3 Oxidative stress .......................................................................................... 370
18.1.4 Nutritional stress ........................................................................................ 370
18.2 Salt tolerance mechanisms in plants...................................................................... 370
18.2.1 Osmotic regulation mechanism ................................................................. 370
18.2.2 Ion homeostasis mechanism ...................................................................... 371
18.2.3 Reactive oxygen species scavenging mechanism ..................................... 371
18.3 Transcription factors involved in salt stress .......................................................... 372
18.3.1 bHLH transcription factors ........................................................................ 372
18.3.2 bZIP transcription factors .......................................................................... 373
18.3.3 NAC transcription factors.......................................................................... 373
xiv Contents

18.3.4 WRKY transcription factors ...................................................................... 374


18.3.5 MYB transcription factors ......................................................................... 374
18.3.6 Other transcription factors participate in salt stress.................................. 375
18.4 Conclusions and perspectives ................................................................................ 376
Acknowledgments .................................................................................................. 377
References............................................................................................................... 377

CHAPTER 19 Plant transcription factors: important factors


controlling oxidative stress in plants .............................................. 383
Shikha Verma, Pankaj Kumar Verma and Debasis Chakrabarty
19.1 Introduction ............................................................................................................ 383
19.2 Oxidative stress and sources .................................................................................. 384
19.2.1 ROS production.......................................................................................... 385
19.3 ROS scavenging ..................................................................................................... 396
19.4 Role of transcription factors in the regulation of stress-responsive genes ........... 397
19.4.1 AP2/ERF family......................................................................................... 398
19.4.2 The bHLH family....................................................................................... 399
19.4.3 MYB family ............................................................................................... 399
19.4.4 The NAC family ........................................................................................ 400
19.4.5 The WRKY family..................................................................................... 400
19.4.6 The bZIP family......................................................................................... 401
19.4.7 The HSF family ......................................................................................... 401
19.5 Conclusion and future prospects............................................................................ 402
Acknowledgments .................................................................................................. 403
References............................................................................................................... 403
Further reading ....................................................................................................... 417

CHAPTER 20 Transcription factors: master regulators of disease


resistance in crop plants.................................................................. 419
Ravi Ranjan Saxesena, Shreenivas Kumar Singh and
Praveen Kumar Verma
20.1 Introduction ............................................................................................................ 419
20.2 Molecular basis of plantmicrobe interaction ...................................................... 420
20.3 The WRKY family of transcription factors and their functional domain ............ 422
20.4 WRKY transcription factors and their role in biotic stress................................... 423
20.5 APETELA2/ethylene-responsive factor family of transcription factors............... 427
20.6 AP2/ERF family of transcription factors and their role in biotic stress ............... 428
20.7 NAC transcription factors and their structural organization ................................. 429
20.8 NAC transcription factors and their role in biotic stress ...................................... 430
Contents xv

20.9 bZIP transcription factors....................................................................................... 433


20.10 bZIP transcription factors and their role in biotic stress....................................... 433
20.11 MYB transcription factor family ........................................................................... 434
20.12 MYB transcription factors and their role in biotic stress...................................... 435
20.13 Conclusion and future perspectives ....................................................................... 435
Acknowledgments .................................................................................................. 436
Declaration of competing interest .......................................................................... 436
Contribution ............................................................................................................ 436
References............................................................................................................... 436

Index .................................................................................................................................................. 445


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List of contributors

Puja Agarwal
Department of Botany, Constituent College, Purnea University, Purnia, Bihar, India
Diksha Bagal
Department of Botany, School of Life Sciences, Central University of Jammu, Samba, Jammu
and Kashmir (UT), India
Lavanya Bhagavatula
Plant Cell and Developmental Biology Laboratory, Department of Biological Sciences, IISER
Bhopal, Bhauri, Madhya Pradesh, India
Alka Bishnoi
Department of Botany, University of Rajasthan, Jaipur, Rajasthan, India
Debasis Chakrabarty
Molecular Biology and Biotechnology Division, CSIR-National Botanical Research Institute,
Lucknow, Uttar Pradesh, India
Divya Chauhan
Division of Germplasm, ICAR-National Bureau of Plant Genetic Resources, New Delhi, Delhi,
India
Rekha Chouhan
CSIR-Indian Institute of Integrative Medicine, Jammu, Jammu and Kashmir, India; Guru Nanak
Dev University (GNDU), Amritsar, Punjab, India
Aksar Ali Chowdhary
Department of Botany, School of Life Sciences, Central University of Jammu, Samba, Jammu
and Kashmir (UT), India
Sourav Datta
Plant Cell and Developmental Biology Laboratory, Department of Biological Sciences, IISER
Bhopal, Bhauri, Madhya Pradesh, India
Vinay Kumar Dhiman
Department of Basic Sciences, Dr. YSP UHF Nauni, Solan, Himachal Pradesh, India
Vivek Kumar Dhiman
Departmnt of Biotechnology, Himachal Pradesh University, Shimla, Himachal Pradesh,
India
Jeremy Dkhar
Plant EvoDevo Laboratory, Agrotechnology Division, CSIR-Institute of Himalayan Bioresource
Technology, Palampur, Himachal Pradesh, India; Academy of Scientific and Innovative
Research (AcSIR), Ghaziabad, Uttar Pradesh, India
Qingkun Dong
Key Laboratory of Plant Molecular Physiology, CAS Center for Excellence in Molecular Plant
Sciences, Institute of Botany, Chinese Academy of Sciences, Beijing, P.R. China

xvii
xviii List of contributors

Sumit G. Gandhi
CSIR-Indian Institute of Integrative Medicine, Jammu, Jammu and Kashmir, India; Academy of
Scientific and Innovative Research (AcSIR), Ghaziabad, Uttar Pradesh, India
Vikrant Gupta
Plant Biotechnology Division, CSIR-Central Institute of Medicinal and Aromatic Plants
(CSIR-CIMAP), Lucknow, Uttar Pradesh, India; Academy of Scientific and Innovative Research
(AcSIR), CSIR-HRDC Campus, Ghaziabad, Uttar Pradesh, India
Gayatri Jamwal
School of Biotechnology, S. K. University of Agricultural Sciences & Technology of Jammu,
Chatha, Jammu and Kashmir, India
Pooja Jangir
Department of Botany, University of Rajasthan, Jaipur, Rajasthan, India
Nikhil Job
Plant Cell and Developmental Biology Laboratory, Department of Biological Sciences, IISER
Bhopal, Bhauri, Madhya Pradesh, India
Zeba Khan
Molecular Biology and Biotechnology Division, CSIR-National Botanical Research Institute,
Lucknow, Uttar Pradesh, India
Ajay Kumar
Molecular Biology and Biotechnology Division, CSIR-National Botanical Research Institute,
Lucknow, Uttar Pradesh, India
Rituraj Kumar
Plant Biotechnology Division, CSIR-Central Institute of Medicinal and Aromatic Plants
(CSIR-CIMAP), Lucknow, Uttar Pradesh, India; Academy of Scientific and Innovative Research
(AcSIR), CSIR-HRDC Campus, Ghaziabad, Uttar Pradesh, India
Isha Magotra
School of Biotechnology, S. K. University of Agricultural Sciences & Technology of Jammu,
Chatha, Jammu and Kashmir, India
Shakti Mehrotra
Department of Biotechnology, Institute of Engineering and Technology, Lucknow, Uttar Pradesh,
India
Sonal Mishra
Department of Botany, School of Life Sciences, Central University of Jammu, Samba, Jammu
and Kashmir (UT), India
Mehali Mitra
Department of Botany, UGC Centre for Advanced Studies, The University of Burdwan, Golapbag
campus, Burdwan, West Bengal, India
Abhilek Kumar Nautiyal
CSIR-Indian Institute of Integrative Medicine, Jammu, Jammu and Kashmir, India; Academy of
Scientific and Innovative Research (AcSIR), Ghaziabad, Uttar Pradesh, India
List of contributors xix

Himanshu Pandey
Department of Biotechnology, Dr. YSP UHF Nauni, Solan, Himachal Pradesh, India
Asosii Paul
Department of Botany, Nagaland University, Lumami, Nagaland, India
Dharitree Phukan
ICAR-National Institute for Plant Biotechnology, New Delhi, Delhi, India
Ujjal Jyoti Phukan
School of Plant Sciences, University of Arizona, Tucson, AZ, United States
Pravin Prakash
Plant Biotechnology Division, CSIR-Central Institute of Medicinal and Aromatic Plants (CSIR-CIMAP),
Lucknow, Uttar Pradesh, India
Puthan Valappil Rahul
Plant Cell and Developmental Biology Laboratory, Department of Biological Sciences, IISER
Bhopal, Bhauri, Madhya Pradesh, India
Garima Rai
CSIR-Indian Institute of Integrative Medicine, Jammu, Jammu and Kashmir, India
Gyanendra K. Rai
School of Biotechnology, S. K. University of Agricultural Sciences & Technology of Jammu,
Chatha, Jammu and Kashmir, India
Hasthi Ram
National Institute of Plant Genome Research, New Delhi, Delhi, India
Nevedha Ravindran
Plant Cell and Developmental Biology Laboratory, Department of Biological Sciences, IISER
Bhopal, Bhauri, Madhya Pradesh, India
Sujit Roy
Department of Botany, UGC Centre for Advanced Studies, The University of Burdwan, Golapbag
campus, Burdwan, West Bengal, India
R.K. Salgotra
School of Biotechnology, S. K. University of Agricultural Sciences & Technology of Jammu,
Chatha, Jammu and Kashmir, India
Ravi Ranjan Saxesena
Plant Immunity Laboratory, National Institute of Plant Genome Research Aruna Asaf Ali Marg,
New Delhi, Delhi, India
Nancy Sharma
CSIR-Indian Institute of Integrative Medicine, Jammu, Jammu and Kashmir, India
Parul Sharma
Biological Central Facility, CSIR-Central Institute of Medicinal and Aromatic Plants, Lucknow,
Uttar Pradesh, India
xx List of contributors

Pooja Kanwar Shekhawat


Department of Botany, University of Rajasthan, Jaipur, Rajasthan, India
Deeksha Singh
Plant Cell and Developmental Biology Laboratory, Department of Biological Sciences, IISER
Bhopal, Bhauri, Madhya Pradesh, India
Devendra Singh
Department of Biotechnology, B.N. College of Engineering and Technology, Lucknow, Uttar
Pradesh, India
Jawahar Singh
Plant Immunity Laboratory, National Institute of Plant Genome Research, Aruna Asaf Ali Marg,
New Delhi, Delhi, India
Shreenivas Kumar Singh
Plant Immunity Laboratory, National Institute of Plant Genome Research Aruna Asaf Ali Marg,
New Delhi, Delhi, India
Vigyasa Singh
Pharmacology and Toxicology Department, College of Pharmacy, University of Arizona, Tucson,
AZ, United States
Praveen Soni
Department of Botany, University of Rajasthan, Jaipur, Rajasthan, India
Rakesh Srivastava
Molecular Biology and Biotechnology Division, CSIR-National Botanical Research Institute,
Lucknow, Uttar Pradesh, India
Vikas Srivastava
Department of Botany, School of Life Sciences, Central University of Jammu, Samba, Jammu
and Kashmir (UT), India
Santosh Kumar Upadhyay
Department of Botany, Panjab University, Chandigarh (UT), India
Pankaj Kumar Verma
Department of Botany, University of Lucknow, Lucknow, Uttar Pradesh, India; French
Associates Institute for Agriculture and Biotechnology of Drylands, Jacob Blaustein Institutes for
Desert Research, Ben-Gurion University of the Negev, Israel
Praveen Chandra Verma
Molecular Biology and Biotechnology Division, CSIR-National Botanical Research Institute,
Lucknow, Uttar Pradesh, India
Praveen Kumar Verma
Plant Immunity Laboratory, School of Life Sciences, Jawaharlal Nehru University, New Delhi,
Delhi, India
List of contributors xxi

Shikha Verma
Molecular Biology and Biotechnology Division, CSIR-National Botanical Research Institute,
Lucknow, Uttar Pradesh, India; French Associates Institute for Agriculture and Biotechnology
of Drylands, Jacob Blaustein Institutes for Desert Research, Ben-Gurion University of the
Negev, Israel
Cui Zhang
Key Laboratory of Plant Molecular Physiology, CAS Center for Excellence in Molecular Plant
Sciences, Institute of Botany, Chinese Academy of Sciences, Beijing, P.R. China; College of Life
Sciences, University of Chinese Academy of Sciences, Beijing, P.R. China
Tingting Zhang
School of Horticulture, Hainan University/Key Laboratory for Quality Regulation of Tropical
Horticultural Crops of Hainan Province, Haikou, P.R. China
Yang Zhou
School of Horticulture, Hainan University/Key Laboratory for Quality Regulation of Tropical
Horticultural Crops of Hainan Province, Haikou, P.R. China
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About the editors

Dr. Vikas Srivastava works as an assistant professor in the Department of


Botany, at the Central University of Jammu in India and currently working
as ‘Royal Society-Newton International Fellow’ at John Innes Centre,
Norwich, United Kingdom. He completed his PhD jointly from the Central
Institute of Medicinal and Aromatic Plants (CSIR-CIMAP) and Lucknow
University, India. Furthermore, he pursued a postdoc at the National
Institute of Plant Genome Research (NIPGR) in New Delhi, India. With
specializations in various aspects of biotechnology, he has acquired the
training and experience to carry out research in diverse fields of plant biol-
ogy. He has published many articles in journals and books of international
repute and worked as a principal investigator for a major project sanc-
tioned by University Grant Commission (UGC) in New Delhi. He is a recipient of various awards
and prestigious fellowships. He delivered several invited lectures for institutes of national repute.

Dr. Sonal Mishra worked as guest faculty in the Department of Botany at


the Central University of Jammu in India. She pursued her PhD jointly from
CSIR-CIMAP in Lucknow and Jawaharlal Nehru University in New Delhi.
Previously, she completed her postdoc as the Dr. D. S. Kothari-Post Doctoral
Fellow (UGC-DSKPDF) and SERB-National Postdoc Fellow (SERB-NPDF)
at the School of Biotechnology, University of Jammu in India. She also
gained postdoc experience at Jawaharlal Nehru University and NIPGR, in
New Delhi, India. With specializations in various aspects of biotechnology,
she has acquired the training and experience to carry out research in diverse
fields of plant biotechnology and molecular biology. She has published many
articles in journals and books of international repute and has received various
awards and prestigious fellowships. She has presented her work in several
seminars and conferences and received appreciation.

Dr. Shakti Mehrotra is a consulting scientist in the Department of


Biotechnology at the Institute of Engineering and Technology, Dr. A.P.J.
Abdul Kalam Technical University in Lucknow, India. She completed her
PhD jointly from CSIR-CIMAP and Lucknow University, India.
Furthermore, she pursued her postdoc at the Institute of Engineering and
Technology, Lucknow (DBT-PDF) and CSIR-CIMAP (DST Young
Scientist). With specializations in various aspects of biotechnology, she
has acquired the training and experience to carry out research in the
diverse fields of plant biotechnology. At present, she is working in the area
of 3D bioprinting and 3D food printing. She has published many articles in
journals and books of national and international repute and worked as a
principal investigator for major projects sanctioned by the Department of

xxiii
xxiv About the editors

Science and Technology (DST) and Department of Biotechnology (DBT), Government of India.
She is a recipient of various awards and prestigious fellowships. She has delivered several presenta-
tions at conferences and workshops and has received appreciation.

Dr. Santosh Kumar Upadhyay currently works as an assistant professor


in the Department of Botany at Panjab University in Chandigarh, India.
Earlier, he was DST-INSPIRE faculty at the National Agri-Food
Biotechnology Institute, Department of Biotechnology, Government of
India in Mohali, Punjab, India. He completed his doctoral work at the
CSIR-National Botanical Research Institute, Lucknow, and was awarded
his PhD in biotechnology from UP Technical University, in Lucknow,
India. He has been working in plant biotechnology for more than 15 years
and currently works in the area of functional genomics. During his doctoral
research, he was involved in the characterization of various insect toxic
proteins such as lectins, chitin-binding proteins, and others from plant bio-
diversity. Currently, his research group at PU has characterized numerous important cation trans-
porters, including calcium cation antiporters, monovalent cation transporters, calcium ATPases, and
important defense-related protein families such as receptor-like kinases, antioxidant enzymes, of
bread wheat. They also characterize long noncoding RNAs related to the abiotic and biotic stress
response. He has also demonstrated the method of genome editing in bread wheat using the
CRISPR-Cas system for functional genomics studies and has developed a tool Sinder for CRISPR
target site prediction. He has authored more than 100 publications, including research papers in
leading journals of international repute, national and international patents, book chapters, and
books. In recognition of his strong credentials and contributions, he has been awarded the NAAS
Young scientist award (201718) and NAAS-Associate (2018) from the National Academy of
Agricultural Sciences, India, INSA Medal for Young Scientist (2013) from the Indian National
Science Academy, India, NASI-Young Scientist Platinum Jubilee Award (2012) from the National
Academy of Sciences, India, and Altech Young Scientist Award (2011). He has also been the recip-
ient of the prestigious DST-INSPIRE Faculty Fellowship (2012), SERB-Early Career Research
Award (2016) from the Ministry of Science and Technology, Government of India, SBS-MKU
Genomics Award (2019) from the Biotech Research Society of India, and several research grants
from various funding agencies like SERB, DBT, and CSIR. Dr. Upadhyay also serves as a member
of the editorial board and a reviewer of many peer-reviewed international journals.
Preface

Transcription factors (TFs) are vital protein molecules involved in biological processes such as
growth, development, and the organism’s response to environmental variables. These protein mole-
cules bind to DNA regulatory sequences and regulate the unique expression of each gene in differ-
ent cell types and during different developmental phases. Analysis and understanding of TF
expression and activities can help researchers establish the significance of their roles in various bio-
logical processes. Scientific exploration and understanding of TFs and their gene regulation is one
of the most dynamic fields of research and technical advancements, giving a wide scope to compile
what is known, what is not known, and how the known facts can be harnessed for future research.
The principal aim of the book Plant Transcription Factors: Contribution in Development,
Metabolism, and Environmental Stress is to provide a resource comprising significant facets of TFs
in plant biology.
The book intends to provide a methodological reserve highlighting several stirring advance-
ments that may strengthen and expand the reader’s knowledge base of the complexity of transcrip-
tional controls of biological processes. The book comprises four sections and includes chapters that
focus on the recent scientific explorations on general aspects of plant TFs and their contribution to
plant development, metabolic processes, and stress management. The chapters compile existing key
concepts as well as advanced information important for better insight into TF targeting and speci-
ficity, the properties of regulatory sequence, and mechanisms of TF action, and present a plant TF
information repertoire. The topics selected are diverse from those included in other methodologi-
cally oriented books on transcriptional regulation of plant biological processes.
Section I of the book includes two chapters (Chapters 1 and 2) that comprise a comprehensive
overview of TFs and their role in plant life. Chapter 1 gives general, yet updated information on
the types of plant TFs. Chapter 2 discusses the role of plant TFs in the adaption of millet in arid
lands.
Section II encompasses Chapters 38, which focus on transcriptional regulation on plant develop-
ment in organogenesis (root, meristem, leaves, fruit, and nodule development) and light-regulated
developments. This section provides current knowledge on the biological functions performed by
various plant TFs and explores the existing molecular data to illustrate how they exert their roles
during plant development.
Section III consists of four chapters (Chapters 912) that comprehensively discuss plant TFs
associated with biochemical changes (biosynthesis and regulation of secondary metabolites such as
alkaloids, flavonoids, and terpenes) and biochemical developments (such as iron homeostasis). This
section sheds light on transcriptional regulation of various biosynthetic pathway genes and signal-
ing cascades to perform highly orchestrated biological functions at the molecular level.
Section IV (Chapters 1320) focuses on plant TFs associated with stress management and miti-
gation that subsequently leads to stress tolerance and survival of plants. Stress, both biotic and abi-
otic, causes severe damage to all plants; however, the extent of damage depends upon the growth
stage and developmental phase of plants. Chapters in this section cover critical discussions and
cross-talks on molecular regulatory strategies of plants, with particular reference to crops to main-
tain vital physiological activities for their survival under various biotic and abiotic stresses during
specific growth stages and developmental phases.

xxv
xxvi Preface

We express our gratitude to all the authors for their outstanding and cutting-edge contributions
to the book. In some instances, this book represents the most current insights, scientific opinions,
and perspectives into the transcriptional regulation of various plant processes, underscoring the
authors’ generosity in sharing their very recent scientific progress in a bench-side reference format.
We would also like to take this opportunity to acknowledge the reviewers of Plant Transcription
Factors: Contribution in Development, Metabolism, and Environmental Stress for their generous
help and suggestions while reviewing chapter manuscripts. We hope this book is a timely contribu-
tion that will serve as an information reserve for the current and next generation of academicians
and scientists working to decipher the mysteries of plant biology by studying the intricate nature of
transcriptional regulation.

Vikas Srivastava
Sonal Mishra
Shakti Mehrotra
Santosh Kumar Upadhyay
SECTION

Plant transcription
factors (TFs) and
general aspects I
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CHAPTER

Plant transcription factors: an


overview of their role in plant life

Aksar Ali Chowdhary1, Sonal Mishra1, Shakti Mehrotra2, Santosh Kumar Upadhyay3,
1
Diksha Bagal1 and Vikas Srivastava1
1
Department of Botany, School of Life Sciences, Central University of Jammu, Samba, Jammu and Kashmir (UT),
India 2Department of Biotechnology, Institute of Engineering and Technology, Lucknow, Uttar Pradesh, India
3
Department of Botany, Panjab University, Chandigarh (UT), India

1.1 Introduction
Transcription factors (TFs) are the master regulator of gene expression and are mainly associated
with plant development, metabolism, and stress management (Yang et al., 2012; Mishra et al.,
2013; Srivastava and Verma, 2015; Srivastava et al., 2017; Baillo et al., 2019). TFs perform gene
regulatory activities by binding to local or distal cis-elements (commonly known as
DNA-binding sites) of genes associated with various biological functions. Once a TF binds on
the DNA binding site of a particular DNA sequence, it brings out the activation or repression
activity through its activator or repressor domains, respectively. TFs may also have other domains
that interact with other proteins, such as other TFs, signaling molecules, etc., which adds to their
functional diversity (Phillips and Hoopes, 2008). For instance, the posttranslational modification
(phosphorylation) of several TFs has been associated with the MAPK cascade system (Guan
et al., 2014). Though TFs are considered the central regulators of the biological functions of plant
life, they are also regulated significantly under diverse internal and external stimuli (Srivastava
and Verma, 2015; Kumar et al., 2016). Further, many plant TFs are known to regulate multiple
functions. For instance, the significance of WRKY and AP2 are reported in stress tolerance and
the regulation of plant-specific metabolism (Mishra et al., 2013, 2015; Kumar et al., 2016). A sig-
nificant portion of the plant genome encodes for TFs, as reported in the case of Arabidopsis,
where .5% of the genes in the genome encodes TFs (Arabidopsis Genome Project; Riechmann
and Ratcliffe, 2000). Hawkins et al. (2021) generated the genome-scale metabolic pathway data-
bases of 126 algal and plant genomes, including model crops and medicinal plants, considering
the significance of TFs. Due to the pivotal role of TFs in the regulation of several molecular
events of plant biology, understanding their functional diversity is immensely important and rele-
vant for exploring the molecular biology of plants. Therefore, this book explores plant TFs and
their functional potential and significance including updated scientific research on TFs.

Plant Transcription Factors. DOI: https://doi.org/10.1016/B978-0-323-90613-5.00003-0


© 2023 Elsevier Inc. All rights reserved. 3
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The main interest, however, of pragmatists in their somewhat
tiresome insistence upon the truism that all truth is made truth is their
hostility (Locke had it in his day) to the supposed rationalist position
that there is an “a priori” and “objective” truth independent altogether
113
of human activities and human purposes. The particular object of
114
their aversion is what Dewey talks of as “that dishonesty, that
insincerity, characteristic of philosophical discussion, that is
manifested in speaking and writing as if certain ultimate abstractions
or concepts could be more real than human purposes and human
beings, and as if there could be any contradiction between truth and
115
purpose.” As we shall reflect at a later stage upon the rationalist
theory of truth, we may, meantime, pass over this hostility with the
remark that it is, after all, only owing to certain peculiar
circumstances (those, say, of its conflict with religion and science
and custom) in the development of philosophy that its first principles
have been regarded by its votaries as the most real of all realities.
These devotees tend to forget in their zeal that the pragmatist way of
looking upon all supposed first principles—that of the consideration
of their utility in and necessity as explanations of our common
experience and its realities—is the only way of explaining their
reality, even as conceptions.
It requires to be added—so much may, indeed, have already
been inferred from the preceding chapter—that, apart from their hint
about the highest truth being necessarily inclusive of the highest
human purposes, it is by no means easy to find out from the
pragmatists what they mean by truth, or how they would define it.
When the matter is pressed home, they generally confess that their
attitude is in the main “psychological” rather than philosophical, that
it is the “making” of truth rather than its “nature” or its “contents” or its
systematic character that interests them. It is the “dynamical” point of
view, as they put it, that is essential to them. And out of the sphere
and the associations of this contention they do not really travel. They
will tell you what it means to hit upon this particular way of looking
upon truth, and how stimulating it is to attempt to do so. And they will
give you many more or less artificial and tentative, external,
descriptions of their philosophy by saying that ideas are “made for
man,” and “not man for ideas,” and so on. But, although they deny
both the common-sense view that truth is a “correspondence” with
external reality, and the rationalist view that truth is a “coherent
system” on its own account, they never define truth any more than
do their opponents the rationalists. It is a “commerce” and not a
116
“correspondence,” they contend, a commerce between certain
parts of our experience and certain other parts, or a commerce
between our ideas and our purposes, but not a commerce with
reality, for the making of truth is itself, in their eyes, the making of
reality.
Secondly, it is another familiar characteristic of Pragmatism that,
although it fails to give a satisfying account either of truth or reality,
the one thing of which it is for ever talking of, as fundamental to our
117
entire life as men, is belief. This is the one thing upon which it
makes everything else to hang—all knowledge and all action and all
theory. And it is, of course, its manifest acceptance of belief as a
fundamental principle of our human life, and as a true measure of
118
reality, that has given to Pragmatism its religious atmosphere. It is
this that has made it such a welcome and such a credible creed to
so many disillusioned and free-thinking people to-day, as well as to
so many of the faithful and the orthodox. “For, in principle,
Pragmatism overcomes the old antithesis of Faith and Reason. It
shows, on the one hand, that faith must underlie all reason and
pervade it, nay, that at bottom rationality itself is the supremest
119
postulate of Faith.” “Truth,” again, as James reminds us, “lives in
fact for the most part on a credit system. Our thoughts and beliefs
[how literally true this is!] pass so long as nobody challenges them,
120
just as bank-notes pass so long as nobody refuses them.”
Now it requires but the reflection of a moment to see that the
various facts and considerations upon which the two last quotations,
and the general devotion of Pragmatism to “belief,” both repose, are
all distinctly in favour of the acceptability of Pragmatism at the
present time. There is nothing in which people in general are more
interested at the beginning of this twentieth century than in belief. It
is this, for example, that explains such a thing as the great success
to-day in our English-speaking world of such an enterprise as the
Hibbert Journal of Philosophy and Religion, or the still greater
phenomenon of the world-wide interest of the hour in the subject of
comparative religion. Most modern men, the writer is inclined to
121
think, believe a great deal more than they know, the chief difficulty
about this fact being that there is no recognized way of expressing it
in our science or in our philosophy, or of acting upon it in our
behaviour in society. It is, however, only the undue prominence of
122
mathematical and physical science since the time of Descartes
that has made evidence and demonstration the main consideration
of philosophy instead of belief, man’s true and fundamental estimate
of reality.
123
We have already pointed out that one of the main results of
Pragmatism is the acceptance on the part of its leading upholders of
our fundamental beliefs about the ultimately real and about the
realization of our most deeply cherished purposes. In fact, reality in
general is for them, we may say—in the absence from their writings
of any better description,—simply that which we can “will,” or
“believe in,” as the basis for action and for conscious “creative”
effort, or constructive effort. As James himself puts it in his book on
The Meaning of Truth: “Since the only realities we can talk about are
objects believed in, the pragmatist, whenever he says ‘reality,’
means in the first instance what may count for the man himself as a
reality, what he believes at the moment to be such. Sometimes the
reality is a concrete sensible presence.... Or his idea may be that of
an abstract relation, say of that between the sides and the
hypotenuse of a triangle.... Each reality verifies and validates its own
idea exclusively; and in each case the verification consists in the
satisfactorily-ending consequences, mental or physical, which the
idea was to set up.”
We shall later have to refer to the absence from Pragmatism of a
criterion for achievement and for “consequences.” And, as far as
philosophical theories are concerned, these are all, to the
pragmatists, true or false simply in so far as they are practically
credible or not. James is quite explicit, for example, about
Pragmatism itself in this regard. “No pragmatist,” he holds, “can
warrant the objective truth of what he says about the universe; he
124
can only believe it.” There is faith, in short, for the pragmatist, in
every act, in every phase of thought, the faith that is implied in the
realization of the purposes that underlie our attempted acts and
thoughts. They eagerly accept, for example, the important doctrine of
the modern logician, and the modern psychologist, as to the
presence of volition in all “affirmation” and “judgment,” seeing that in
every case of affirmation there is a more or less active readjustment
of our minds (or our bodies) to what either stimulates or impedes our
activity.
A third outstanding characteristic of Pragmatism is the “deeper”
view of human nature upon which, in contrast to Rationalism, it
supposes itself to rest, and which it seeks to vindicate. It is this
supposedly deeper view of human nature for which it is confessedly
pleading when it insists, as it is fond of doing, upon the connexion of
philosophy with the various theoretical and practical pursuits of
mankind, with sciences like biology and psychology, and with social
125
reform, and so on. We have, it may be remembered, already
intimated that even in practical America men have had their doubts
about the depth of a philosophy that looks upon man as made in the
main for action and achievement instead of, let us say, the
realization of his higher nature. Still, few of the readers of James can
have altogether failed to appreciate the significance of some of the
many eloquent and suggestive paragraphs he has written upon the
limitations of the rationalistic “temperament” and of its unblushing
sacrifice of the entire wealth of human nature and of the various
pulsating interests of men to the imaginary exigencies of abstract
126
logic and “system.” To him and to his colleagues (as to Socrates,
for that part of it) man is firstly a being who has habits and purposes,
and who can, to some extent, control the various forces of his nature
through true knowledge, and in this very discrepancy between the
real and the ideal does there lie for the pragmatists the entire
problem of philosophy—the problem of Plato, that of the attainment
of true virtue through true knowledge.
Deferring, however, the question of the success of the
pragmatists in this matter of the unfolding of the true relation
between philosophy and human nature, let us think of a few of the
teachings of experience upon this truly important and inevitable
relation, which no philosophy indeed can for one moment afford to
neglect. Insistence upon these facts or teachings and upon the
reflections and criticisms to which they naturally give rise is certainly
a deeply marked characteristic of Pragmatism.
Man, as has often been pointed out, is endowed with the power
of reflection, not so much to enable him to understand the world
either as a whole or in its detailed workings as to assist him in the
further evolution of his life. His beliefs and choices and his spiritual
culture are all, as it were, forces and influences in this direction.
Indeed, it is always the soul or the life principle that is the important
thing in any individual or any people, so far as a place in the world
(or in “history”) is concerned.
Philosophers, as well as other men, often exchange (in the
words of Lecky) the “love of truth” as such for the love of “the truth,”
that is to say, for the love of the system and the social arrangements
that best suit their interests as thinkers. And they too are just as
eager as other men for discipleship and influence and honour.
Knowledge with them, in other words, means, as Bacon put it,
“control”; and even with them it does not, and cannot, remain at the
stage of mere cognition. It becomes in the end a conviction or a
belief. And thus the philosopher with his system (even a Plato, or a
Hegel) is after all but a part of the universe, to be judged as such,
along with other lives and other systems—a circumstance hit off
early in the nineteenth century by German students when they used
to talk of one’s being able (in Berlin) to see the Welt-Geist (Hegel)
“taking a walk” in the Thiergarten.
Reality again, so far as either life or science is concerned,
means for every man that in which he is most fundamentally
interested—ions and radium to the physicist of the hour, life to the
biologist, God to the theologian, progress to the philanthropist, and
so on.
Further, mankind in general is not likely to abandon its habit of
estimating all systems of thought and philosophy from the point of
view of their value as keys, or aids, to the problem of the meaning
and the development of life as a whole. There is no abstract “truth”
or “good” or “beauty” apart from the lives of beings who contemplate,
and who seek to create, such things as truth and goodness and
beauty.
To understand knowledge and intellect, again, we must indeed
look at them in their actual development in connexion with the total
vital or personal activity either of the average or even of the
exceptional individual. And instead of regarding the affections and
the emotions as inimical to knowledge, or as secondary and inferior
to it, we ought to remember that they rest in general upon a broader
and deeper attitude to reality than does either the perception of the
127
senses or the critical analysis of the understanding. In both of
these cases is the knowledge that we attain to limited in the main
either to what is before us under the conditions of time and space, or
to particular aspects of things that we mark off, or separate, from the
totality of things. As Bergson reminds us, we “desire” and “will” with
the “whole” of our past, but “think” only with “part” of it. Small wonder
then that James seeks to connect such a broad phenomenon as
religion with many of the unconscious factors (they are not all merely
“biological”) in the depth of our personality. Some of the instincts and
the phenomena that we encounter there are things that transcend
altogether the world that is within the scope of our senses or the
reasoning faculties.
Truth, too, grows from age to age, and is simply the formulated
knowledge humanity has of itself and its environment. And errors
disappear, not so much in consequence of their logical refutation, as
in consequence of their inutility and of their inability to control the life
and thought of the free man. Readers of Schopenhauer will
remember his frequent insistence upon this point of the gradual
dissidence and disappearance of error, in place of its summary
refutation.
Our “reactions” upon reality are certainly part of what we mean
by “reality,” and our philosophy is only too truly “the history of our
heart and life” as well as that of our intellectual activity. The historian
of philosophy invariably acts upon a recognition of the personal and
the national and the epochal influence in the evolution of every
philosophical system. And even the new, or the fuller conception of
life to which a given genius may attain at some stage or other of
human civilization will still inevitably, in its turn, give place to a newer
or a more perfect system.
Now Pragmatism is doubtless at fault in seeking to create the
impression that Rationalism would seek to deny any, or all, of those
characteristic facts of human nature. Still, it is to some extent
justified in insisting upon their importance in view of the sharp
conflict (we shall later refer to it) that is often supposed to exist
between the theoretical and the practical interests of mankind, and
that Rationalism sometimes seems to accept with comparative
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equanimity. What Pragmatism is itself most of all seeking after is
a view of human nature, and of things generally, in which the fullest
129
justice is done to the facts upon which this very real conflict of
modern times may be said to rest.
A fourth characteristic of Pragmatism is its notorious “anti-
130
intellectualism,” its hostility to the merely dialectical use of terms
131
and concepts and categories, to argumentation that is unduly
detached from the facts and the needs of our concrete human
experience. This anti-intellectualism we prefer meantime to consider
not so much in itself and on its own account (if this be possible with a
negative creed) as in the light of the results it has had upon
philosophy. There is, for example, the general clearing of the ground
that has undoubtedly taken place as to the actual or the possible
meaning of many terms or conceptions that have long been current
with the transcendentalists, such as “pure thought,” the “Absolute,”
“truth” in and for itself, philosophy as the “completely rational”
interpretation of experience, and so on. And along with this clearing
of the ground there are (and also in consequence of the pragmatist
movement) a great many recent, striking concessions of Rationalism
to practical, and to common-sense, ways of looking at things, the
very existence of which cannot but have an important effect upon the
philosophy of the near future. Among some of the more typical of
these are the following:
From Mr. F. H. Bradley we have the emphatic declarations that
the principle of dialectical opposition or the principle of “Non-
Contradiction” (formerly, to himself and his followers, the “rule of the
game” in philosophy) “does not settle anything about the nature of
reality”; that “truth” is an “hypothesis,” and that “except as a means
to a foreign end it is useless and impossible”; and “when we judge
truth by its own standard it is defective because it fails to include all
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the facts,” and because its contents “cannot be made intelligible
throughout and entirely”; that “no truth is idle,” and that “all truth” has
“practical” and æsthetic “consequences”; that there is “no such
133
existing thing as pure thought”; that we cannot separate truth and
practice; that “absolute certainty is not requisite for working
134
purposes”; that it is a “superstition to think that the intellect is the
highest part of us,” and that it is well to attack a one-sided
“intellectualism”; that both “intellectualism” and “voluntarism” are
“one-sided,” and that he has no “objection to identifying reality with
goodness or satisfaction, so long as this does not mean merely
135
practical satisfaction.” Then from this same author comes the
following familiar statement about philosophy as a whole:
“Philosophy always will be hard, and what it promises in the end is
no clear vision nor any complete understanding or vision, but its
certain reward is a continual and a heightened appreciation [this is
the result of science as well as of philosophy] of the ineffable
mystery of life, of life in all its complexities and all its unity and all its
136
worth.”
Equally typical and equally important is the following concession
from Professor Taylor, although, of course, to many people it would
seem no concession at all, but rather the mere statement of a fact,
which our Neo-Hegelians have only made themselves ridiculous by
seeming to have so long overlooked: “Mere truth for the intellect can
never be quite the same as ultimate reality. For in mere truth we get
reality only in its intellectual aspect, as that which affords a higher
satisfaction to thought’s demand for consistency and systematic
unity in its object. And as we have seen, this demand can never be
137
quite satisfied by thought itself. For thought, to remain thought,
must always be something less than the whole reality which it
138
knows.”
And we may add also from Professor Taylor the following
declaration in respect of the notorious inability of Neo-Hegelian
Rationalism to furnish the average man with a theory of reality in the
contemplation of which he can find at least an adequate motive to
conscious effort and achievement: “Quite apart from the facts, due to
personal shortcomings and confusions, it is inherent in the nature of
metaphysical study that it can make no positive addition to our
information, and can itself supply no motive for practical
139
endeavour.”
Many of those findings are obviously so harmonious with some
of the more familiar formulas of the pragmatists that there would
seem to be ample warrant for associating them with the results of the
pragmatist movement. This is particularly the case, it would seem,
with the concession of Mr. Bradley with respect of the “practical” or
“hypothetical” conception that we ought to entertain of “truth” and
“thinking,” and also with the strictures passed by him upon “mere
truth” and “mere intellectualism,” and with Professor Taylor’s position
in respect of the inadequacy of the rationalist theory of reality, as in
no sense a “dynamic” or an “incentive” for action. And we might well
regard Professor Taylor’s finding in respect of mere systematic truth
or the “Absolute” (for they are the same thing to him) as confirmatory
of Dr. Schiller’s important contention that “in Absolutism” the two
“poles” of the “moral” and the “intellectual” character of the Deity “fall
apart.” This means, we will remember, that the truth of abstract
140
intellectualism is not the truth for action, that absolutism is not
able to effect or harmonize between the truth of systematic
knowledge and moral truth—if, indeed, there be any such thing as
moral truth on the basis of a pure Rationalism.
To be sure, both the extent and even the reality of all this
supposed cession of ground in philosophy to the pragmatists has
been doubted and denied by the representatives of Rationalism.
They would be questioned, too, by many sober thinkers and scholars
who have long regarded Hegelian intellectualism and pragmatist
“voluntarism” as extremes in philosophy, as inimical, both of them, to
the interests of a true and catholic conception of philosophy. The
latter, as we know from Aristotle, should be inclusive of the realities
both of the intellectual and the practical life.
Pragmatist criticisms of Rationalism, again, may fairly be claimed
to have been to a large extent anticipated by the independent
findings of living idealist thinkers like Professors Pringle-Pattison,
Baillie, Jones, and others, in respect of the supposed extreme claims
of Hegelianism, as well as by similar findings and independent
constructive efforts on the part of the recent group of the Oxford
141
Personal Idealists. That there is still a place for pragmatist anti-
intellectualism is evidently the conclusion to be drawn from such
things as the present wide acceptance of the philosophy of Bergson,
or the recent declarations of Mr. Bradley that we are justified “in the
intelligent refusal to accept as final an theoretical criterion which
actually so far exists,” and that the “action of narrow consistency
must be definitely given up.”
The reflection ought, moreover, to be inserted here that even if
Pragmatism has been of some possible service in bringing forth from
rationalists some of their many recent confessions of the limitations
of an abstract intellectualism, it is not at all unlikely that Rationalism
in its turn may succeed in convicting Pragmatism of an undue
142
emphasis upon volition and action and upon merely practical
truth.
We shall now terminate the foregoing characterization of
Pragmatism by a reference to two or three other specific things for
which it may, with more or less justice, be supposed to stand in
philosophy. These are (1) the repudiation of the “correspondence
143
view” of the relation of truth to reality, (2) the rejection of the idea
of there being any ultimate or rigid distinction between “appearance”
and “reality,” and (3) the reaffirmation of the “teleological” point of
view as characteristic of philosophy in distinction from science.
As for (1) it has already been pointed out that this idea of the
misleading character of the ordinary “correspondence notion” of truth
is claimed by pragmatists as an important result of their proposal to
test truth by the standard of the consequences involved in its
144
acceptance. The ordinary reader may not, to be sure, be aware of
the many difficulties that are apt to arise in philosophy from an
apparent acceptance of the common-sense notion of truth as
somehow simply a duplicate or a “copy” of external reality. There is
the difficulty, say, of our ever being able to prove such a
correspondence without being (or “going”) somehow beyond both
the truth and the reality in question, so as to be able to detect either
coincidence or discrepancy. Or, we might again require some bridge
between the ideas in our minds and the supposed reality outside
them—“sensations” say, or “experiences,” something, in other words,
that would be accepted as “given” and indubitable both by idealists
and realists. And there would be the difficulty, too, of saying whether
we have to begin for the purposes of all reflective study with what is
within consciousness or with what is outside it—in matter say, or in
things. And if the former, how we can ever get to the latter, and vice
versa. And so on with the many kindred subtleties that have divided
thinkers into idealists and realists and conceptualists, monists,
dualists, parallelists, and so on.
Now Pragmatism certainly does well in proposing to steer clear
of all such difficulties and pitfalls of the ordinary “correspondence
notion.” And as we shall immediately refer to its own working
philosophy in the matter, we shall meantime pass over this mere
point of its rejection of the “correspondence notion” with one or two
remarks of a critical nature, (1) Unfortunately for the pragmatists the
rejection of the correspondence notion is just as important a feature
145
of Idealism as it is of Pragmatism. The latter system therefore can
lay no claim to any uniqueness or superiority in this connexion. (2)
Pragmatism, as we may perhaps see, cannot maintain its position
that the distinction between “idea” and “object” is one “within
experience itself” (rather than a distinction between experience and
something supposedly outside it) without travelling further in the
146
direction of Idealism than it has hitherto been prepared to do. By
such a travelling in the direction of Idealism we mean a far more
thorough-going recognition of the part played in the making of reality
by the “personal” factor, than it has as yet contemplated either in its
“instrumentalism” or in its “radical empiricism.” (3) There is, after all,
an element of truth in the correspondence notion to which
Pragmatism fails to do justice. We shall refer to this failure in a
147
subsequent chapter when again looking into its theory of truth
and reality.
Despite these objections there is, however, at least one
particular respect in regard to which Pragmatism may legitimately
claim some credit for its rejection of the correspondence notion. This
is its insistence that the truth is not (as it must be on the
correspondence theory) a “datum” or a “presentation,” not something
given to us by the various objects and things without us, or by their
supposed effects upon our senses and our memory and our
understanding. It rather, on the contrary, maintains Pragmatism, a
“construction” on the part of the mind, an attitude of our “expectant”
(or “believing”) consciousness, into which our own reactions upon
things enter at least as much as do their supposed effects and
impressions upon us. Of course the many difficulties of this thorny
subject are by no means cleared up by this mere indication of the
148
attitude of Pragmatism, and we shall return in a later chapter to
this idea of truth as a construction of the mind instead of a datum,
taking care at the same time, however, to refer to the failure of which
we have spoken on the part of Pragmatism to recognize the element
of truth that is still contained in the correspondence notion.
(2) The rejection of the idea of any rigid, or ultimate distinction
between “appearance” and “reality.” This is a still broader rejection
than the one to which we have just referred, and may, therefore, be
thought of as another more or less fundamental reason for the
rejection either of the copy or of the correspondence theory of truth.
The reality of things, as Pragmatism conceives it, is not something
already “fixed” and “determined,” but rather, something that is
“plastic” and “modifiable,” something that is, in fact, undergoing a
continuous process of modification, or development, of one kind or
another. It must always, therefore, the pragmatist would hold, be
defined in terms of the experiences and the activities through which
it is known and revealed and through which it is, to some extent,
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even modified.
Pragmatism, as we may remember, has been called by James
“immediate” or “radical” empiricism, although in one of his last books
he seeks to give an independent development to these two
doctrines. The cardinal principle of this philosophy is that “things are
what they are experienced as being, or that to give a just account of
150
anything is to tell what that thing is experienced to be.” And it is
perhaps this aspect of the new philosophy of Pragmatism that is
most amply and most attractively exhibited in the books of James. It
is presented, too, with much freshness and skill in Professor
151
Bawden’s book upon Pragmatism, which is an attempt, he says,
“to set forth the necessary assumptions of a philosophy in which
152
experience becomes self-conscious as a method.”
153
“The new philosophy,” proceeds Bawden, “is a pragmatic
idealism. Its method is at once intrinsic and immanent and organic or
functional. By saying that its method is functional, we mean that its
experience must be interpreted from within. We cannot jump out of
our skins ... we cannot pull ourselves up by our own bootstraps. We
find ourselves in mid-stream of the Niagara of experience, and may
define what it is by working back and forth within the current.” “We
do not know where we are going, but we are on the way” [the
contradiction is surely apparent]. Then, like James, Bawden goes on
to interpret Pragmatism by showing what things like self-
consciousness, experience, science, social consciousness, space,
time, and causation are by showing how they “appear,” and how they
“function”—“experience” itself being simply, to him and to his friends,
a “dynamic system,” “self-sustaining,” a “whole leaning on nothing.”
The extremes of this “immediate” or “radical” philosophy appear
to non-pragmatists to be reached when we read words like those just
quoted about the Niagara stream of our experience, and about our
life as simply movement and acceleration, or about the celebrated “I
think” of Descartes as equally well [!] set forth under the form “It
thinks,” or “thinking is going on,” or about the “being” of the individual
person as consisting simply in a “doing.” “All this we hold,” says
Bawden, “to be not materialism but simply energism.” “There is no
‘truth,’ only ‘truths’—this is another way of putting it—and the only
criterion of truth is the changing one of the image or the idea which
comes out of our impulses or of the conflict of our habits.” The end of
all this modern flowing philosophy is, of course, the “Pluralism” of
James, the universe as a society of functioning selves in which
reality “may exist in a distributive form, or in the shape, not of an All,
but of a set of eaches.” “The essence of life,” as he puts it in his
154
famous essay on Bergson, “is its continually changing character,”
and we only call it a “confusion” sometimes because we have grown
accustomed in our sciences and philosophies to isolate “elements”
155
and “differents” which in reality are “all dissolved in one another.”
“Relations of every sort, of time, space, difference, likeness, change,
rate, cause, or what not, are just as integral members of the
sensational flux as terms are.” “Pluralism lets things really exist in
the each form, or distributively. Its type of union ... is different from
the monistic type of all-einheit. It is what I call the strung-along type,
the type of continuity, contiguity, or concatenation.” And so on.
(3) The reaffirmation of the teleological point of view. After the
many illustrations and references that have already been given in
respect of the tendencies of Pragmatism, it is perhaps hardly
necessary to point out that an insistence upon the necessity to
philosophy of the “teleological” point of view, of the consideration of
both thoughts and things from the point of view of their purpose or
utility, is a deeply-marked characteristic of Pragmatism. In itself this
demand can hardly be thought of as altogether new, for the idea of
considering the nature of anything in the light of its final purpose or
end is really as old in our European thought as the philosophy of
Aristotle or Anaxagoras. Almost equally familiar is the kindred idea
upon which Pragmatism is inclined to felicitate itself, of finding the
roots of metaphysic “in ethics,” in the facts of conduct, in the facts of
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the “ideal” or the “personal” order which we tend in human
civilization to impose upon what is otherwise thought of by science
as the natural order. The form, however, of the teleological argument
to which Pragmatism may legitimately be thought to have directed
our attention is that of the possible place in the world of reality, and
in the world of thought, of the effort and the free initiative of the
individual. This place, unfortunately (the case is quite different with
157
Bergson ), Pragmatism has been able, up to the present time, to
define, in the main, only negatively—by means of its polemic against
the completed and the self-completing “Absolute” of the Neo-
Hegelian Rationalists. What this polemic is we can best indicate by
quoting from Hegel himself a passage or a line of the reflection
against which it is seeking to enter an emphatic and a reasoned
protest, and then after this a passage or two from some of our Anglo-
Hegelians in the same connexion.
“The consummation,” says Hegel, in a familiar and often-quoted
passage, “of the Infinite aim (i.e. of the purpose of God as
omniscient and almighty) consists merely in removing the illusion
158
which makes it seem unaccomplished.” Now although there is a
sense in which this great saying must for ever be maintained to
159
contain an element of profound truth, the attitude of Pragmatism
in regard to it would be, firstly, that of a rooted objection to its
outspoken intellectualism. How can the chief work of the Almighty be
conceived to be merely that of getting rid somehow from our minds,
or from his, of our mental confusions? And then, secondly, an equally
rooted objection is taken to the implication that the individual human
being should allow himself to entertain, as possibly true, a view of
the general trend of things that renders any notion of his playing an
160
appreciable part therein a theoretical and a practical absurdity.
This notion (or “conceit,” if you will) he can surrender only by ceasing
to think of his own consciousness of “effort” and of the part played by
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“effort” and “invention” in the entire animal and human world, and
also of his consciousness of duty and of the ideal in general. This
latter consciousness of itself bids him to realize certain “norms” or
regulative prescripts simply because they are consonant with that
higher will which is to him the very truth of his own nature. He
cannot, in other words, believe that he is consciously obliged to work
and to realize his higher nature for nothing. The accomplishment of
ends and of the right must, in other words, be rationally believed by
him to be part of the nature of things. It is this conviction, we feel
sure, that animates Pragmatism in the opposition it shares both with
common sense and with the radical thought of our time against the
162
meaninglessness to Hegelianism, or to Absolutism, many of the
hopes and many of the convictions that we feel to be so necessary
and so real in the life of mankind generally.
And there are other lines of reflection among Neo-Hegelians
against which Pragmatism is equally determined to make a more or
less definite protest, in the interest, as before, of our practical and of
our moral activity. We may recall, to begin with, the memorable
words of Mr. Bradley, in his would-be refutation of the charge that the
ideals of Absolutism “to some people” fail to “satisfy our nature’s
demands.” “Am I,” he indignantly asks, “to understand that we are to
have all we want, and have it just as we want it?” adding (almost in
the next line) that he “understands,” of course, that the “views” of
Absolutism, or those of any other philosophy, are to be compared
“only with views” that aim at “theoretical consistency” and not with
163
mere practical beliefs. Now, speaking for the moment for
Pragmatism, can it be truly philosophical to contemplate with
equanimity the idea of any such ultimate conflict as is implied in
164
these words between the demands of the intellect and the
demands of emotion—to use the term most definitely expressive of a
personal, as distinct from a merely intellectual satisfaction?
Then again there is, for example, the dictum of Dr. McTaggart,
that there is “no reason to trust God’s goodness without a
demonstration which removes the matter from the sphere of
165
faith.” May there not, we would ask, be a view of things according
to the truth of which the confidence of the dying Socrates in the
reasonableness and the goodness of God are at least as reasonable
as his confession, at the same time, of his ignorance of the precise,
or the particular, fate both of the just and of the unjust? And is not,
too, such a position as that expressed in these words of Dr.
McTaggart’s about a logically complete reason for believing in the
essential righteousness of things now ruled out of court by some of
the concessions of his brother rationalists to Pragmatism, to which
reference has already been made? It is so ruled out, for example,
even by Mr. Bradley’s condemnation as a “pernicious prejudice” of
the idea that “what is wanted for working purpose is the last
166
theoretical certainty about things.”
CHAPTER IV
PRAGMATISM AND HUMAN ACTIVITY

It requires now but a slight degree of penetration to see that beneath


this entire matter of an apparent opposition between our “theoretical”
and our “practical” satisfaction, and beneath much of the pragmatist
insistence upon the “consequences” of ideas and of systems of
thought, there is the great question of the simple fact of human
action and of its significance for philosophy. And it might truly be said
that the raising of this question is not merely another of the more or
less definitely marked features of Pragmatism, but in some respects
it is one outstanding characteristic.
For some reason or other, or for some strange combination of
167
reasons, the phenomenon that we call “action” (the activity of
man as an agent) and the apparently simple facts of the reality and
the intelligibility of action have long been regarded as matters of
altogether secondary or subordinate importance by the rationalism of
philosophy and by the mechanical philosophy of science. This
Rationalism and this ostensibly certain and demonstrable
mechanical philosophy of science suppose that the one problem of
human thought is simply that of the nature of truth or of the nature of
reality (the reality of the “physical” world) as if either (or each) of
these things were an entity on its own account, an absolutely final
finding or consideration. That this has really been the case so far as
philosophy is concerned is proved by the fact even of the existence
of the many characteristic deliverances and concessions of
Rationalism in respect of Pragmatism to which reference has already
been made in the preceding chapter. And that it has also been the
case so far as science is concerned is proved by the existence of the
many dogmatic attempts of many natural philosophers from Holbach
to Haeckel to apply the “iron laws” of matter and motion to the reality
168
of everything else under heaven, and of everything in the
heavens in spite of the frequent confessions of their own colleagues
with regard to the actual and the necessary limits and limitations of
science and of the scientific outlook.
Only slowly and gradually, as it were, has the consideration
come into the very forefront of our speculative horizon that there is
for man as a thinking being no rigid separation between theory and
practice, between intellect and volition, between action and thought,
169
between fact and act, between truth and reality. There is clearly
volition or aim, for example, in the search after truth. And there is
170
certainly purpose in the attention that is involved even in the
simplest piece of perception, the selection of what interests and
affects us out of the total field of vision or experience. And it is
equally certain that there is thought in action—so long, that is to say,
as action is regarded as action and not as impulse. Again, the man
who wills the truth submits himself to an imperative just as surely as
does the man who explicitly obeys the law of duty. It is thus
impossible, as it were, even in the so-called intellectual life, to
distinguish absolutely between theoretical and practical
considerations—“truth” meaning invariably the relations obtaining in
some “sphere,” or order, of fact which we separate off for some
purpose or other from the infinite whole of reality. Equally impossible
is it to distinguish absolutely between the theoretical and the
practical in the case of the highest theoretical activity, in the case,
say, of the “contemplation” that Aristotle talks of as the most
“godlike” activity of man. This very contemplation, as our Neo-
171
Hegelian friends are always reminding us, is an activity that is just
as much a characteristic of man, as is his power of setting his limbs
in motion.
We have referred to the desire of the pragmatists to represent,
and to discover, a supposedly deeper or more comprehensive view
of human nature than that implicitly acted upon by Intellectualism—a
view that should provide, as they think, for the organic unity of our
active and our so-called reflective tendencies. This desire is surely
eminently typical of what we would like to think of as the rediscovery
by Pragmatism for philosophy, of the active, or the volitional, aspects
of the conscious life of man, and along with this important side of our
human nature, the reality also of the activities and the purposes that
are revealed in what we sometimes speak of as unconscious nature.
The world we know, it would hold, in the spirit and almost in the letter
172
of Bergson, lives and grows by experiment, and by activities and
processes and adjustments. Pragmatism has doubtless, as we
pointed out, been prone to think of itself as the only philosophy that
can bake bread, that can speak to man in terms of the actual life of
effort and struggle that he seems called upon to live in the
environment in which he finds himself. And, as we have just been
insisting, the main ground of its hostility to Rationalism is the
apparent tendency of the latter to treat the various concepts and
hypotheses that have been devised to explain the world, and to
render it intelligible, as if they were themselves of more importance
than the real persons and the real happenings that constitute the
173
world of our experience.
If it were at all desirable to recapitulate to any extent those
phenomena connected with Pragmatism that seem to indicate its
rediscovery of the fact of action, and of the fact of its meaning for
philosophy, as its one outstanding characteristic, we may point to
such considerations as the following: (1) The fact of its having
sought to advance from the stage of a mere “instrumentalist” view of
human thought to that of an outspoken “humanism” or a socialized
utilitarianism. (2) The fact of its seeking to leave us (as the outcome
of philosophy) with all our more important “beliefs,” with a general
“working” view of the world in which such things as religion and
ideals and enthusiasm are adequately recognized. Pragmatism is
really, as we have put it, more interested in belief than in knowledge,
the former being to it the characteristic, the conquering attitude of
man to the world in which he finds himself. (3) Its main object is to
establish a dynamical view of reality, as that which is “everywhere in
the making,” as that which signifies to every person firstly that aspect
of the life of things in which he is for the time being most vitally

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