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Environmental Plant
Physiology
Taylor & Francis
Taylor & Francis Group
http://taylorandfrancis.com
Environmental Plant
Physiology

Neil Willey

GS Garland Science
Taylor & Francis Group
NEW YORK LONDON
Vice President: Denise Schanck
Assistant Editor: David Borrowdale
Production Editor: Georgina Lucas
Illustrators: Neil Willey and Nigel Orme
Layout: Georgina Lucas
Cover Designer: Andrew Magee
Copy Editor: Josephine Hargreaves
Proofreader: Sally Livitt
Indexer: Bill Johncocks

© 2016 by Garland Science, Taylor & Francis Group, LLC

Neil Willey is Reader in Environmental Plant Physiology at the University of the West of England
(UWE), Bristol, UK. He holds a BSc (Hons) in Biology & Geography and a PhD in Botany, both from the
University of Bristol, UK. He’s an active teacher of plant biology to undergraduate and postgraduate
students from a variety of disciplines. His research focuses on the behavior and effects of pollutants,
especially radioisotopes, in the soil-plant system. He’s the Director of the UWE Graduate School and the
Chair of the UK Coordinating Group for Environmental Radioactivity.

Front Cover.
Close-up of Rhododendron flower.

This book contains information obtained from authentic and highly regarded sources. Every effort has
been made to trace copyright holders and to obtain their permission for the use of copyright material.
Reprinted material is quoted with permission, and sources are indicated. A wide variety of references
are listed. Reasonable efforts have been made to publish reliable data and information, but the author
and the publisher cannot assume responsibility for the validity of all materials or for the consequences
of their use. All rights reserved. No part of this publication may be reproduced, stored in a retrieval
system or transmitted in any form or by any means—graphic, electronic, or mechanical, including
photocopying, recording, taping, or information storage and retrieval systems—without permission of
the copyright holder.

ISBN 978-0-8153-4469-8

Library of Congress Cataloging-in-Publication Data

Names: Willey, Neil, author..
Title: Environmental plant physiology / Neil Willey.
Description: New York, NY : Garland Science, 2016.
Identifiers: LCCN 2015039662 | ISBN 9780815344698 (alk. paper)
Subjects: LCSH: Plant ecophysiology.
Classification: LCC QK717 .W55 2016 | DDC 571.2--dc23
LC record available at http://lccn.loc.gov/2015039662

Published by Garland Science, Taylor & Francis Group, LLC, an informa business,
711 Third Avenue, New York, NY, 10017, USA, and 3 Park Square, Milton Park, Abingdon, OX14 4RN, UK.

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Preface

Environmental plant physiology focuses on the foundations of life on land,

with direct implications for the possibility of humans inhabiting Earth
sustainably. I hope that readers of this book will gain an understanding of
the importance of plant–environment interactions and be inspired to help
humanity face some of its major challenges—in particular, global food
security and the conservation of the natural world. This book is intended
for upper-level undergraduate and graduate students, but will also be use-
ful to some researchers. Several excellent textbooks inspired my interest
in plant–environment interactions, but so significant are recent scientific
advances that I felt a new textbook was needed. Its approach is intended to
be useful across the biological, environmental, and agricultural sciences. In
an era in which science is progressively focusing on the major challenges
facing humanity, the ability to engage with a topic using both disciplinary
and trans-disciplinary perspectives is increasingly important. In this book
my aim is to provide a synthesis that is useful to individual biological, envi-
ronmental, and agricultural disciplines, whilst also providing a framework
for understanding across them all.
As is evident from the structure of the book, I think that a useful contem-
porary understanding of environmental plant physiology can be built by
focusing on environmental variables, each of which is important to the bio-
logical, environmental, and agricultural sciences. These environmental vari-
ables are grouped into ‘resources’, ‘stressors’, and ‘xenobiotics’ according to
whether they help synthesize biomass, limit biomass production, or poison
plants. Each chapter is structured to focus on plant responses to progres-
sively more profound variation in the environmental variable. This is used to
suggest a hierarchy of responses from molecular to ecological scales in each
chapter, and then as an overall framework in the final chapter. Throughout,
examples are drawn from both unmanaged and managed ecosystems, and
the importance of evolutionary history is acknowledged whenever possible
to help to explain the occurrence of adaptations.
In each chapter the text is enhanced by large boxes that include additional
stories of interest, small boxes in the margins that elucidate some key points,
and by tables and figures. In particular, I have made an effort to provide a new
set of figures for understanding environmental plant physiology. Further
read­ing and references for each section are listed at the end of each chapter,
and all bold terms are defined in the glossary. The book is complemented by
resources available to students online: the glossary and flashcards; image
gallery of the key species discussed in the text as indicated in bold green font;
audio files that provide a commentary for each topic; and a set of multiple-
choice questions. For instructors, all of the figures from the book are available
to download in both PowerPoint® and JPEG format.
Overall, this textbook provides a heuristic framework that enables students,
in the midst of the torrent of information published in scientific journals, not
only to think about guiding principles and concepts in environmental plant
physiology, but also to form their own perspectives. I would be interested in
any feedback from anyone (contact me at Neil.Willey@uwe.ac.uk).
vi PREFACE

Acknowledgments
I am very grateful to the students who have taken my courses; the discus-
sion that their interest stimulates has been vital to developing this book. I
would also like to express my appreciation for the very helpful comments of
colleagues and reviewers. Many thanks to staff from Garland Science: Dave
Borrowdale for steering the project from beginning to end, Gina Almond for
her role in getting it off the ground, and Georgina Lucas for all her efforts
during the publication process. And finally, particular thanks must go to
Lorna and the rest of my family without whose support this book, and much
else, would never have happened.
Neil Willey

Reviewers
Habib-ur-Rehman Athar (Bahauddin Zakariya University); Juergen
Burkhardt (University of Bonn); Ivan Couée (University of Rennes); Peter
S. Curtis (The Ohio State University); Stephen Ebbs (Southern Illinois
University); David E. Evans (Oxford Brookes University); Ros Gleadow
(Monash University); Kevin Griffin (Columbia University); Stuart Lane
(Plymouth University); Richard Leegood (University of Sheffield); Denis
Murphy (University of South Wales); Bob Nowak (University of Nevada,
Reno); Léon-Etienne Parent (Université Laval); David Tissue (University
of Western Sydney); Alyson Tobin (University of St Andrews); Marcos
Yanniccari (CONICET, Universidad Nacional de La Plata); Jianhua Zhu
(University of Maryland).
 vii

Contents
Chapter 1 Contexts, Perspectives, Leaf optical properties are adapted to long-term
and Principles 1 variation in light regimes 41
Plant interactions with the atmosphere, hydrosphere, Adjustments in leaf position and plant architecture
and geosphere underpin terrestrial ecosystems 1 adapt plants to different light regimes 44
Minimizing human impact on ecosystems and Photoinhibition is most severe in alpine
achieving global food security are significant environments 46
challenges 3 Summary 48
Proximate and ultimate questions elucidate how and Further reading 49
why plants interact with the environment 5
Resources, stressors, and toxins affect plant biomass
production and quality 6 Chapter 3 Carbon Dioxide 51
Environmental factors that affect plant growth are CO2 fixation underpins the primary production of
interacting but independent variables 10 biomass 51
Many reference soil groups are a product of Variation in the supply of CO2 to plants is significant
interacting environmental variables 10 and affected by human activity 53
Spatial and temporal analyses provide insights into The regulation of rubisco activity controls CO2 entry
plant responses to environmental variation 11 into the Calvin–Benson cycle 56
Plants process information about environmental Oxygenation of RuBP decreases growth but provides
variation using signaling networks 14 rapid metabolic flexibility 58
Differences in gene expression and in the genes When there is a sustained low CO2 supply, C4
expressed underpin a hierarchy of plant adaptations 14 plants maintain a high CO2:O2 ratio in the vicinity
of rubisco 60
Environmental plant physiology is ecologically useful
in defining plant traits and niches 19 C3–C4 intermediates and C4 plants show distinct
responses to chronic differences in the
Studying plant–environment interactions can help to environment 63
increase agricultural efficiency and sustainability 20
Crassulacean acid metabolism adapts plants to
Modeling is improving our understanding of plant– chronically difficult CO2-fixation conditions 66
environment interactions 21
Long-term increased CO2 levels can increase
Summary 21 plant growth, but limiting factors can moderate
Further reading 22 this effect 69
Plant responses to increasing CO2 levels will affect
the hydrological cycle and Earth’s climate 72
Chapter 2 Light 23 An understanding of CO2 fixation by plants is
In plants, ancient photosynthetic systems provide important for sustainable food production and
the chemical energy for terrestrial ecosystems 23 ecosystem conservation 73
Photosystems, cytochromes, and ATP synthases Summary 75
transduce light energy into chemical energy 25 Further reading 75
Terrestrial plants have to adapt to a generally high
and very variable light regime 28
Plants can adjust quickly to variation in PAR using Chapter 4 Water 77
non-photochemical quenching 31 Plant–water relations affect physiological processes
Plants can adjust electron flows to help them to from a cellular to a global scale 77
withstand variable light intensities 34 Water management is vital for ensuring global food
PSII repair is important in plants that tolerate high security and minimizing the impact of human
light intensities 36 activity on the environment 80
Chloroplast movements can be used to adjust fairly Water potential gradients drive water movement,
rapidly the amount of light absorbed 37 including transpiration in trees over 100 m tall 83
Photosystems, grana, and thylakoids adapt to Short-term adjustments of resistance to water flux
differences in light regime 39 allow water homeostasis 85
viii CONTENTS

Many plants adapt physiologically to short-term Mycorrhizas are major adaptations for phosphorus
water deficit 88 acquisition in low-phosphorus environments 142
Extended water deficit induces changes in Some species use cluster root systems to
root growth 90 intensively mine phosphorus from the soil 146
Leaf adaptations aid drought survival and provide Carnivorous plants digest organic phosphorus
alternative ways of capturing water 92 using phosphatases 150
Succulent xerophytes are physiologically Summary 150
decoupled from their chronically arid environments 94 Further reading 151
Resurrection plants cope with complete desiccation 95
Interactions between water and other stressors
provide important environmental insights 99 Chapter 7 Essential and
Summary 100 Beneficial Elements 153
Further reading 101 Terrestrial plants evolved to mine the soil for an
ancient suite of available elements 153
The availability of essential nutrients limits
Chapter 5 Nitrogen 103 biomass production and quality in many
Nitrogen assimilated in plants is vital for the ecosystems 156
production of biomolecules in terrestrial organisms 103 Elemental homeostasis is achieved using both
Artificially fixed nitrogen significantly affects the ion-binding compounds and transport proteins 157
biosphere and atmosphere 104 Plants adjust to a variable supply of
The concentration of different forms of soil micronutrients by overexpressing homeostatic
nitrogen varies significantly 107 components 159
Plant nitrogen-transporter uptake capacity is Beneficial elements help many plant species to
tuned to variation in soil nitrogen supply 111 cope with a wide range of abiotic stresses 160
Sub-optimal sulfur availability can inhibit the
Plants integrate nitrogen from different sources by
synthesis of ecophysiologically important
converting it to NH3 for assimilation 113
compounds 162
Whole-plant physiological adjustments help to use
Potassium can limit ecosystem production, but
different patterns of nitrogen supply 115
its use in fertilizer has a moderate environmental
Plants adjust their root morphology in response to impact 164
shortages of nitrogen 116
Calcium deficiency can occur in a variety of
Symbioses contribute significantly to plant plants, and magnesium deficiency in a variety
nitrogen uptake in nitrogen-deficient environments 117 of crops 166
Carnivorous plants are mixotrophs that can Adaptations of root anatomy and morphology
obtain nitrogen opportunistically from an erratic help plants to respond to chronic nutrient
supply 123 deficiency 168
Summary 126 Many plants use symbioses with fungi and
Further reading 127 changes in rhizosphere microflora to aid nutrient
uptake 170
Ionomics 171
Chapter 6 Phosphorus 129 Summary 173
Phosphorus availability often controls terrestrial Further reading 174
biomass production and ecosystem processes 129
Current phosphorus fertilizer regimes are
unsustainable, inefficient, and often polluting 131 Chapter 8 Temperature 175
Phosphorus homeostasis is a key challenge for Plants are static poikilotherms, so significant
plants in terrestrial ecosystems 133 variation in temperature is a considerable
Plants have numerous transporters that regulate challenge 175
uptake and translocation 135 Changing global temperature regimes are affecting
Plants can increase the availability of inorganic plant growth, development, and distribution 177
phosphorus and the breakdown of organic Plants detect temperature changes via physical
phosphorus 136 changes in numerous biomolecules 180
Plants can adjust their root system morphology Chilling, freezing, and heat initiate changes in key
to optimize phosphorus uptake 140 components of different signaling pathways 183
 CONTENTS ix

In some plants, chilling temperatures can induce Plant cells have multiple mechanisms for buffering
an acclimation response based on the CBF regulon 184 cytosolic pH 233
Adaptation to non-optimal temperature Acid soils contain high solution concentrations
necessitates maintaining membranes in the of ions that are toxic to plant cells 234
liquid–crystal state 186 Some plants resist the effects of moderate soil
Freezing-tolerant plants produce cryoprotectants acidity by excluding aluminum from the
and osmoprotectants 188 cytoplasm 237
Heat-tolerant plants have protein curation For many plants on acid soils, mycorrhizal
mechanisms adapted to increase the rate of associations increase aluminum resistance 240
protein repair 191 On very acidic soils, some plants take up and
Anatomical and morphological adaptations of compartmentalize aluminum 241
leaves aid plant tolerance of prolonged cold Basic soils are low in important nutrients and
and heat 194 induce characteristic symptoms in plants 243
Temperature-induced physiological changes trigger Some plants have adapted to scavenge iron, zinc,
developmental and phenological responses 198 and manganese from basic soils 246
Summary 199 Nicotianamine aids iron homeostasis, and in
Further reading 199 grasses evolved into root exudates that chelate iron 247
Ecologically important iron and zinc deficiency
responses are finding important agricultural uses 249
Chapter 9 Salinity 201 Summary 250
Terrestrial plants are descended from freshwater
Further reading 251
algae, so saline water is generally toxic to them 201
Plant responses to salinity are important in
irrigated agriculture and in salt marshes and Chapter 11 Flooding 253
mangrove swamps 204
Flooding is a significant variable in both
Exposure to salt induces osmotic and ionic unmanaged and managed terrestrial ecosystems 253
stresses in plants 208
Human activity is adversely affecting wetlands
Sodium can enter plants via symplastic and and increasing the incidence of flooding 255
apoplastic pathways, but can be removed from
Waterlogged soils are low in oxygen and some
the cytoplasm 211
nutrients, but high in toxins 255
Salt-tolerant plants compartmentalize sodium,
Soil waterlogging rapidly induces hypoxia,
and halophytes also control potassium:sodium
cellular acidosis, and decreased water uptake 258
ratios 213
Physiological adjustments enable some plants to
At high salinity, halophytes synthesize
withstand soil waterlogging for short periods 259
specialized metabolites in order to adapt to
osmotic challenges 215 Ethylene signaling is central to plant responses to
excess water 261
Salt tolerance in crops has been increased by
manipulating biochemical and physiological In many plants, waterlogging-induced hypoxia
traits 217 induces changes in root anatomy 262
Halophytes that face severe osmotic stresses have Wetland plants form extensive constitutive
morphological and physiological adaptations 219 aerenchyma and adapt morphologically to
flooding 266
Some halophytes use specialized organs to
excrete sodium chloride from their leaves 221 In some flooded soils, pneumatophores help
woody plants to aerate their roots 268
Mangrove and salt-marsh plants tolerate
waterlogging and salinity 223 The adaptations of wetland plants often produce
oxidized rhizospheres 269
Summary 224
Some plants can adapt to submergence of their
Further reading 225
shoots 271
Emergent aquatic macrophytes can force oxygen
Chapter 10 Soil pH 227 down through organs buried deep in anoxic mud 273
Soil pH affects the growth of both wild and Some aquatic macrophytes are adapted to living
domesticated plants 227 permanently submerged 275
Soil pH is operationally defined and human Summary 275
activities are affecting it on a global scale 229 Further reading 276
x CONTENTS

Chapter 12 Inorganic Toxins 279 Summary 329

A few reactive elements are essential, but they Further reading 329
and many non-essential elements can also
be toxic 279
Human activity is significantly increasing the Chapter 14 Air Pollutants 331
concentrations of inorganic toxins in the Earth’s Plants are dependent on an extensive surface area
ecosystems 281 that interacts with the atmosphere 331
Homeostatic mechanisms control the uptake and Adverse effects of air pollution on plants will be
translocation of reactive elements in plants 284 important in the twenty-first century 334
Exposure to inorganic toxins decreases growth The deposition of air pollutants on plants depends
and reproduction via physiological and genetic on the properties of plants and pollutants 336
effects 288 Plants can assimilate some sulfur dioxide, but
Amplified homeostatic mechanisms in the roots anthropogenic deposition rates can exceed this
of some species produce a metal-tolerant capacity 337
physiology 292 Direct uptake of gaseous reactive nitrogen species
Some plants have the capacity to minimize the can affect plant growth and ecosystem dynamics 339
uptake of toxins from high external concentrations 295 Semi-volatile and volatile organic compounds
Some plants can hyperaccumulate inorganic can be absorbed by and released from vegetation 342
toxins in their shoots 296 Chronic effects of ozone on terrestrial plants
Chronic exposure to toxins in metalliferous will be significant in the twenty-first century 345
ecosystems provides some unique biological Particulates filtered by plants from the
insights 300 atmosphere can affect their growth 348
Control of soil-to-plant transfer of inorganic Plants can be used to monitor and manage air
toxins is useful in agriculture and quality 350
phytoremediation 301
Summary 351
Summary 303
Further reading 352
Further reading 303

Chapter 15 Synopsis and Outlook 355

Chapter 13 Organic Toxins 305
Plant–environment interactions play a significant
Plants can control the reactivity of many organic role in determining the boundaries of non-linear
functional groups 305 effects in Earth systems 355
Synthetic organic compounds underpin modern The understanding of plant stress response
life but can have a significant environmental mechanisms can be extended by comparisons
impact 308 with other organisms 356
The entry of organic toxins into plants depends Our understanding of the importance of
on soil, plant, and chemical properties 309 variation in plant–environment interactions can
Organic toxins elicit reactive and perhaps also be extended by modeling that includes the pattern
proactive stress responses in plants 313 and scale of variation 358
In plant cells, many organic toxins can be Understanding how plant stress responses evolved
transformed enzymatically 315 will provide insights about the plant–environment
In some plants, organic toxins and their interface 359
transformation products can be deactivated by Understanding plant–environment interactions
conjugation 317 helps us to confront global challenges 361
In some plants, conjugated organic toxins can be Further reading 362
allocated to metabolically inactive compartments 319
Non-target-site herbicide resistance can evolve
from xenobiotic detoxification mechanisms 320 Abbreviations list 363
Target-site resistance helps plants to adapt to Glossary 365
catastrophic exposure to herbicides 323
Plants enhance the bioremediation of water and Index 378
soils contaminated with organic xenobiotics 324
Manipulation of plant tolerance of organic toxins
is of increasing importance 327
 light
air CO2
pollutants

organic water
toxins

Chapter 1
Contexts,
inorganic nitrogen
toxins

Perspectives, flooding phosphorus

and Principles soil pH

essentials
and
beneficials

salinity temperature

Key concepts
• Plant–environment interactions are the foundation of terrestrial Plant growth is affected by variation
ecosystems. in independent, interacting
environmental variables.
• Environmental change and food security are significant challenges for
humankind.
• Physiology is the study of how and why organisms function as they do.
• Biomass production and quality are dependent on resources, stressors,
and xenobiotics.
• The environmental factors that affect plants are independent variables
with overlapping effects.
• Major soil types embody the effects of many interacting variables that
influence plant growth.
• Spatial and temporal variation provides key perspectives on plant–
environment interactions.
• Plants detect environmental variation via signal transduction networks.
• There is a hierarchy of adaptations that underpin plant responses to
environmental variation.
• Environmental plant physiology can elucidate key ecological processes.
• Agricultural systems can benefit from insights into plant–environment
interactions.
• Models can be used to predict plant behavior in a changing
environment.

Plant interactions with the atmosphere,

hydrosphere, and geosphere underpin terrestrial
ecosystems
The colonization of the land surface by multicellular plants was a momen-
tous phase in the history of life on Earth, in significant part because multicell­
ular plants in effect colonized the atmosphere, providing an unprecedented
link between the subsurface, surface, and above-surface environments.
It initiated perhaps the most significant ever increase in the number of
niches and ecosystems. The evolution of organisms into these new niches
2 CHAPTER 1 Contexts, Perspectives, and Principles

transformed not only the domains of life but also the biogeochemistry of
Earth. Terrestrial plants are therefore at a crucial nexus of the biogeochemi-
cal cycles of the Earth, and help to provide the life-support system for terres-
trial species, including humans. Understanding plant function at this nexus
provides major insights into many of the environmental challenges that face
humankind. This book aims to provide an understanding of plant physiol-
ogy that is informed by the development of terrestrial ecosystems and rele­
vant to current environmental challenges.

The earliest evidence of multicellular plants that were adapted to the chal-
lenges of living on land is provided by spore tetrad microfossils from the
Ordovician period. These suggest that, in some terrestrial locations at least,
there were quite extensive stands of plants on land by 450 million years ago.
Due to the lack of macrofossils, it is uncertain what these plants looked like,
but they were probably liverwort-like and inhabited wet environments,
perhaps living in shallow standing water. Macrofossils from the subsequent
Silurian period suggest that, by 425 million years ago, plants on land were
up to 10 cm tall and had rhizoids—they were beginning to function partly
in the atmosphere and partly in the regolith. Between 425 and 300 million
years ago there was a profound increase in the diversity of terrestrial plants
(Figure 1.1). Complex terrestrial ecosystems began to develop in which a
diverse range of plants, many of which have descendants in current ecosys-
tems, adapted to the challenges of life on land. Fossils from the Devonian
and Carboniferous periods show that some of these plants were many tens
of meters tall, and although many of them were clearly swamp dwellers,
some probably inhabited drier habitats.

Detailed understanding of the environmental physiology of early plants is dif-

ficult, but numerous features of extant terrestrial plants that are interpreted
as adaptations to the challenges of life on land were evident in some of the
early terrestrial plants. These include adaptations for gas exchange with the
atmosphere, for water transport, and for nutrient uptake. It was the profound
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Figure 1.1. The evolution of major
terrestrial plant phyla. Extant major 200
groups are shown, but not the first Triassic
3.7 billion years of Earth’s history 251
(pre-Cambrian), nor many fossil and Permian
minor extant groups. Characteristics in 300
ovules/seeds
green letters are major adaptations that
Carboniferous
affected plant–environment relationships.
PALEOZOIC

megaphylls
Mycorrhizae = root symbioses with fungi; 375
microphylls = small simple leaves without Devonian
extensive vascular systems; megaphylls = 420
vascular systems, root systems, mycorrhizae, microphylls
Silurian
large true leaves with extensive vascular 455
systems; ovules = structures that contain Ordovician
spores, cuticles, stomata
the female gametophyte. (Redrawn from 490
Ridge I [2002] Plants. With permission Cambrian
from Oxford University Press.) 542
 Minimizing human impact on ecosystems and achieving global food security are significant challenges 3

environmental changes which these adaptations eventually wrought that

affected the biogeochemistry of Earth—for example, the composition of the
atmosphere from this time on was affected by the activity of terrestrial plants ATMOSPHERE
and the decomposition of their dead biomass. The cycling of fresh water was
transformed so that a significant proportion of all the water moving from
the land to the atmosphere did so through plants. Following the injection
of organic matter into the regolith, true soils became extensive for the first BIOSPHERE GEOSPHERE
time, transforming the geochemistry of the land surface. Terrestrial plants
are therefore integral to terrestrial ecosystems at perhaps the most impor-
tant interface between the biosphere and the atmosphere, hydrosphere, and
geosphere (Figure 1.2). HYDROSPHERE

An ecosystem can be defined as a community of organisms and the physi-

cal environment with which it interacts, via flows of energy and nutrients,
to develop trophic relationships. This book focuses on the interaction of
multicellular terrestrial plants with their abiotic environment—that is, one Figure 1.2. Plants at a primary
facet of terrestrial ecosystems. It does so in a way that is useful not only to interface. The soil is a primary interface
between the atmosphere, hydrosphere,
those whose particular interest in terrestrial ecosystems relates to the plant
and geosphere. Plants function at this
and biological sciences, but also to those whose primary interests are in the interface, affecting many of the most
environmental and agricultural sciences. To achieve this, a range of ecosys- important biogeochemical cycles on Earth,
tems are discussed in terms of “unmanaged” and “managed” ecosystems, to and providing the foundation of terrestrial
reflect some of the systematic differences between “natural” and “agricul- ecosystems.
tural” terrestrial ecosystems. It is indisputable that, in addition to the top-
ics covered here, understanding of both plant interactions with the biotic
components of the environment and of ecosystem functioning will be vital
to meeting the environmental challenges that face humankind.

Minimizing human impact on ecosystems and

achieving global food security are significant
challenges
If, as many geologists posit, humans are now a primary agent of envi-
ronmental change, the current geological epoch can be defined as the
Anthropocene. There is much evidence that hunting by early humans had
significant adverse impacts on the megafauna of numerous regions on
Earth—for example, large mammals were often hunted to extinction after
the arrival of humans. However, it was with the initiation of agriculture that
humans began to have effects on the environment that were detectable on a
wide scale. The invention of agriculture between 10,000 and 8000 years ago
is perhaps the most significant event in the human story, and it has been
suggested that it defines the start of the Anthropocene. However, despite the
environmental impact of early humans, it is generally accepted that in the
twentieth century the human impact on terrestrial ecosystems was unprec-
edented and initiated “something new under the sun”—that is, a truly global
scale of environmental impact on Earth by a single species. The Millennium
Ecosystem Assessments, the United Nations Environment Program Global
Environmental Outlook (“GEO”) reports, and numerous other assessments
have described the global scale of human impact on terrestrial ecosystems,
which now constitutes one of the greatest challenges facing humankind.
The social and natural sciences will both clearly have vital roles in meeting
this challenge. A striking number of aspects of the latter and many of the
potential solutions do, however, occur at the nexus of plant–environment
interactions. Some of the most important impacts of changing CO2 concen-
trations are, directly or indirectly, on plant growth, but plants also help to
control atmospheric CO2 concentrations. Threats to unmanaged ecosys-
tems include uncontrolled exploitation by humans for food, fuel, and plant
products, and contamination with nutrients and xenobiotics, but manipula-
tion of plant growth might also help to solve some of these problems. Such
current challenges and potential solutions mean that in order to minimize
4 CHAPTER 1 Contexts, Perspectives, and Principles

human impacts on the environment, an understanding of the interactions of

plants with their abiotic environment is probably more important now than
it has ever been.

In the 1970s it was difficult to predict when global population growth might
stop, but now most credible predictions suggest that the global popula-
tion will peak at 10–12 billion in the second half of the twenty-first century.
This is a very significant increase in population when the strain on terres-
trial ecosystems from the current population of about 7 billion is already
so significant. The intensification of agriculture during the latter part of the
twentieth century was one of the most profound and successful of the many
applications of science that were developed in that century. In 1900 it was
inconceivable that it would be possible to come anywhere close to meeting
current demand for agricultural products used for food, forage, fuel, cloth-
ing, drinks, or raw materials. A modest proportion of the increase in agri-
cultural productivity in the twentieth century was due to an increase in the
total area of agricultural production, but the greatest proportion was due to
an increase in the amount of resource per unit area per unit time being con-
verted into yield. This was often achieved by adding resources and by using
varieties that matured more quickly to enable production of more than one
crop per year. Thomas Malthus famously suggested in 1798 that population
growth would inevitably outstrip agricultural production, but the history of
agriculture suggests that humans are adept at avoiding Malthus’s prediction.
They have done so in significant part by increasing the efficiency of plant–
environment interactions in agricultural ecosystems.

The intensification of agriculture in the twentieth century, and in particular

the Green Revolution in developing agriculture, probably saved hundreds
of millions of lives. The food, forage, and plant products that its techniques
produce now underpin, to a remarkable extent, twenty-first-century societ-
ies. In significant part because of its great success, intensive agriculture is
now intimately connected to many of the world’s environmental challenges.
These include demand for water, nutrients, and agrochemicals, together with
the problems that their production and use entail. It has been recognized
since the later years of the twentieth century that in order to achieve global
food security in the twenty-first century, while at the same time reducing the
environmental impact of agriculture, there is a necessity for a “doubly green”
revolution—that is, an increase in production similar to that of the Green
Revolution, together with the minimization of agriculture’s environmental
impact. If this occurs, it is likely to involve a profound change in the way
that plants are used to convert inorganic resources from the environment
into food, forage, and plant products, and it is likely to be dependent on an
appreciation of plant–environment interactions in both unmanaged and
managed ecosystems. For example, a very significant pressure on agriculture
is the scarcity of land on which to grow crops. This arises in part because of
the restricted range of plants that are used to provide the majority of food. It
has been long been recognized that many species of plants other than those
currently used might be useful for producing food, and that natural ecosys-
tems often have highly efficient resource-use patterns. The great variety of
species that are used to produce forage and plant products is a reminder of
the diversity that is potentially available for harvest. The history of alterna-
tive crops, agroforestry, intercropping, and numerous other agricultural
systems runs deeper than is often supposed, and in the twenty-first century
such systems might play a significant role in helping humans to harvest food
and other plant products sustainably. If this happens, an understanding of
plant–environment interactions is likely to aid their development. In this
book, the minimizing of human impact on the environment, along with the
sustainable harvesting of food, forage, and other plant products, are used as
key contexts for understanding plant–environment interactions.
 Proximate and ultimate questions elucidate how and why plants interact with the environment 5

Proximate and ultimate questions elucidate how

and why plants interact with the environment
Physiology is the study of function in living systems. Function is most fully
understood by asking both proximate and ultimate questions. Molecular
biology transformed the plant sciences, and it is now possible to describe
in unprecedented detail the changes that occur in plants in response to a
change in the environment. We are now more able than ever before to answer
questions such as “How do plants respond to a decrease in soil pH?” This
book provides an overview of how plants respond to variation in key envi-
ronmental variables. The ability to describe the mechanistic details of how
something happens—that is, its proximate cause—is vital to understanding
plant–environment interactions. Understanding plant function necessitates
investigations of proximate causes on a scale ranging from that of biomol-
ecule to that of whole organism.

However, it is also possible to ask “Why do plants respond as they do to

decreasing temperature?” Such questions are aimed at the ultimate cause of
a phenomenon (Box 1.1), and in this book are regarded as integral to physi-
ology as the study of function. Multicellular terrestrial plants have an evo-
lutionary history. DNA sequences have an imprint of past events, via both
genetic and epigenetic mechanisms, which legitimizes the asking of ulti-
mate questions. Thus, for example, plants respond as they do to decreases
in temperature in significant part because one of the most significant cold
periods in the last 250 million years on Earth occurred very recently, ending
only 10,000 years ago. In this book, plant physiology is defined as the study
of “how and why a plant functions as it does.”

Phylogenetics has transformed our understanding of the evolutionary his-

tory of many groups of organisms, including plants (Box 1.2), providing
unprecedented answers to ultimate questions. Evolutionary relationships
between many terrestrial plants are now known, providing a phylogenetic
perspective on function. For example, despite the diversity and advantages
of nitrogen-fixing symbioses, phylogenetics shows—perhaps surprisingly—
that the propensity to form these symbioses in flowering plants evolved in
only one group. The likely phylogeny of the angiosperms that constitute the
majority of plant biomass on Earth and many of the groups mentioned in
this book is shown in Figure 1.3. The perspectives derived from proximate
and ultimate questions, informed by molecular biology and phylogenetics,

BOX 1.1. PROXIMATE AND ULTIMATE QUESTIONS

ABOUT PLANT BEHAVIOR

The study of animal behavior is regarded as a discipline in its own right,

namely ethology. Plants also react, respond, and adapt to their environment.
They do this to enhance their chances of survival by, for example, foraging for
resources, avoiding stress, and communicating between individuals. This is,
in a very real sense, “plant behavior.” Ethologists often distinguish between
proximate and ultimate questions about behavior. These are “Tinbergen’s
questions,” and they have echoes of some of the “causes” of animal character-
istics distinguished by Aristotle. The biologist Ernst Mayr was the most prom-
inent proponent of the utility of proximate and ultimate causes in explaining
biological phenomena. Proximate questions focus on the mechanisms that
explain how a behavior or phenomenon is produced within the lifespan of an
individual. Ultimate questions focus on explaining why the particular behav-
ior or phenomenon occurs in the species.
6 CHAPTER 1 Contexts, Perspectives, and Principles

BOX 1.2. MOLECULAR PHYLOGENETICS

Centuries of study have identified many of the major groups
of plants, so if the evolution of a particular character is being
investigated, an outgroup, which is highly likely to be dis-
tantly related, can be chosen to root an analysis. Using an out-
group enables a rooted evolutionary tree to be constructed,
rather than a simple cluster of similarity. A major complica-
tion in phylogenetic reconstruction is long-branch attraction,
in which a small change in a character can produce a radical
reorganization of the long branches of a phylogenetic tree. The
use of algorithms to control for long-branch attraction and of
multiple characters increases the likelihood of identifying the
evolutionary pathways that actually occurred. When variance
in a molecular character such as a gene sequence is known,
Monte Carlo methods can be used to produce what are in
effect infinitely large data sets from which numerous phy-
logenetic trees can be reconstructed. Consensus trees can
then be used to estimate the probability of each node having
occurred. Measures such as bootstrap and jackknife values
quantify how often particular nodes occur in reconstructed
consensus trees, and thus estimate their likelihood of having
occurred (Figure 1).

1 2 3 4 5 6 7 8 9 10 11

95

Figure 1. Contrasting possible phylogenies. The black lines

A C
for A and B show two diametrically opposite possible branching
patterns that could have given rise to 11 taxa. Mostly evolutionary
pathways have occurred somewhere in between (green lines for C),
but the number of possible trees is very large. The likelihood of each
node having occurred can be estimated statistically (for example,
B jackknife percentage shown on inset).

respectively, are used in each chapter of this book to elucidate plant function.
The discussion of function in each chapter starts with molecular biology and
proceeds towards function of whole plants in a phylogenetic context.

Resources, stressors, and toxins affect plant

biomass production and quality
Plants, as autotrophs, utilize inorganic resources from the environment to
synthesize the organic molecules necessary for life. The fundamental chal-
lenge facing terrestrial plants involves gathering the resources necessary to
drive autotrophy. The plants of terrestrial ecosystems are almost all photoau-
totrophs, so light is their fundamental resource. Light energy is used to syn-
thesize organic compounds based primarily on carbon (C), hydrogen (H),
oxygen (O), nitrogen (N), sulfur (S), and phosphorus (P) atoms. These com-
pounds include the carbohydrates, amino acids, and lipids that are essential
requirements for almost all other organisms in terrestrial ecosystems. This
book focuses first on plant capture of resources (Figure 1.4), because of their
importance for all inhabitants of terrestrial ecosystems, including humans.
Resources are defined by their necessity for the production of the biomol-
ecules that are essential for life, and by the adverse effects that a sub-optimal
supply can have. The use in ecology of the concepts underpinning “the cap-
ture of resources,” with their implications of competition for and exploit­
ation of resources, developed synergistically with their use in economics.
However, they have provided a powerful perspective for understanding how
18 CHAPTER 1 Contexts, Perspectives, and Principles

Figure 1.8. The inhibition of gene

expression by small RNAs. (1) In plant MIR CH3
NUCLEUS
genomes, in addition to protein-coding ARGO
genes there are numerous DNA sequences dsDNA NRPD1b
that code for double-stranded microRNAs pri-miRNA DCL
(miRNAs). These miRNA sequences are (5) gene expression
(1)
palindromic, producing hairpin primary DCL
miRNA (pri-miRNA) structures about 70 DCL pre-miRNA mRNA
nucleotides long. These are processed, in
part using Dicer-like proteins (DCLs) that
can slice RNA, to produce double-stranded miRNA (3)
miRNAs about 19–23 nucleotides long
that are exported to the cytoplasm. (2) In DCL
the cytoplasm these bind to the RNA- (2)
mRNA
induced silencing complex (RISC), which DCL ARGO
RISC
includes ARGONAUT proteins (ARGO) ARGO
with RNA-binding and slicing capacity. mRNA
(3) ARGO-bound RNA binds to mRNA
from gene expression, slicing it to prevent
(4)
its translation in ribosomes. In plants,
RdRP ARGO gene silencing
miRNA sequences can be very specific to 3ʹ ssRNA
UTRs, inhibiting the translation of specific
genes, although some miRNAs can inhibit ssRNA
translation of several genes. (4) It is likely ribosome
that miRNA inhibition of mRNA translation defense against
viruses
evolved from plant defense against the
many single-stranded RNA (ssRNA) viruses
that attack plants. ARGO can slice ssRNAs,
or RNA-dependent RNA polymerases
(RdRPs) convert ssRNA virus sequences The microRNAs are made from RNAs about 70 nucleotides long that are
into double-stranded RNA (dsRNA) that
can be attacked by DCLs and attached to
encoded in the genome. These have palindromic sequences towards their
the RISC complex for slicing. (5) The RISC ends, form hairpin loops, and are then cleaved by Dicer-like enzymes to
complex also has a role in methylating form miRNAs. A ribonuclease complex, the RNA-induced silencing com-
genes to exert epigenetic control over their plex, uses the miRNAs to attach to specific mRNAs and cleave them, initi-
expression. ating a cascade of RNA degradation that prevents translation (Figure 1.8).
This mechanism of preventing mRNA translation probably evolved from
defense against the single-stranded RNA viruses common in plants, and is
also the
EPP basis
Figure the short-interfering
for 1.8 RNAs (siRNAs)
inhibition of expression that RNAs
by small can be
P used to
silence the expression of specific genes.

If translation is initiated, polypeptides begin to emerge from ribosomes.

The conformation of most proteins is essential to their function, but can-
not be finalized until translation is complete. There are therefore numer-
ous mechanisms in plants that control protein folding and assembly. These
can be important in the regulation of translation into a proteome response
to particular stressors. In plants, low-intensity or short-term responses can
induce significant changes in gene expression. The physiology of a plant at
a given moment in time is therefore in part a product of the expression of its
genes and the condition of its environment. However, there are also mech­
anisms through which the conditions that a plant or a population of plants
has experienced in the past can affect gene expression and adaptation to the
environment. Epigenetic differences in gene expression are heritable but do
not arise from differences in gene sequence. The methylation of the cyto-
sine and adenine bases in DNA, and the acetylation of the histones around
which DNA is wrapped, and which can be altered by environmental con-
ditions, can result in heritable differences in gene expression. To combat
intense or sustained environmental variation, a combination of constitutive
differences in gene expression and differences in the genes expressed also
occurs, and can give rise to the different anatomies, morphologies, and phe-
nologies that adapt plants to their environment. In general, anatomical and
morphological plant adaptations to intense or chronic differences in envi-
ronmental variables are complementary to biochemical and physiological
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be clarified, and with which the birds must be covered, when cold, to
the thickness of at least half an inch in each pot. These will be
excellent, taken cold.

russian polony.
Your success in this undertaking chiefly depends upon the choice
of the meats made use of. Take of
The lean of Belfast smoked hams, 1 year old 3 lb.
The fat of Belfast smoked hams, 1 year old 3 lb.
High flavoured hung beef 3 lb.
Smoked ox tongues 3 lb.
Hard back fat of bacon 4 lb.
Peel the tongues after being boiled and reject the roots and tips. Cut
the four first of the above up into dice and pound them separately
into smooth pastes, with mucilage of gum tragacanth and fresh
butter. Next cut the fat bacon into dice or cubes as large as the finest
growth of peas, no rind or gristles to be retained; then mix intimately
together,
Garlic, minced 1½ oz.
Shalots, minced 6 oz.
Juniper berries, in fine powder 3 oz
Jamaica pepper, in fine powder 3 oz.
Black peppercorns 3 oz.
Bay salt, in fine powder 1 lb.
Coarse sugar ¾ lb.
and blending all the meats well, season highly with the mixture, and
put it down in a jar for a week to mellow. Then work the mass well for
half an hour. Get some of the largest ox intestines, and soak them in
luke-warm salt and water, wipe them dry and proceed to fill them,
keeping out the air as much as possible, which you will effect by
regular and even pressure, and pricking, only where necessary, with
a stocking-needle. Make your polonies about a foot long each, and
put them aside as done until the next day; then repeat the pressure
and tie them up finally. They must be put into boiling-water with a
little salt and saltpetre, and after once boiling, simmer only for half an
hour. Then take them up, wipe dry, and hang in a current of air for a
week, being turned daily without fail. Then smoke them with
Oak lops 2 parts
Beech chips 2 parts
Fern 1 part
Peat 1 part
for a month. Stow them away in malt cooms. They will be prized by
those persons who delight in high flavours. They need no coating;
nay, many foreigners prefer them mouldy on the outsides.

german saveloys.
Take rough Hambro’ smoked beef 2 lb.
Neats’ tongues,
smoked 1 lb.
Smoked ham 1 lb.
Fat of bacon 2 lb.
Boil the tongue moderately, peal and cut off the gross root tip. Cut up
the whole of the meat into large dice, and pound it thoroughly, mixing
the fat in equally in all parts of the mass. Pick out all skins, sinews,
&c., and mix a pound of good moist sugar throughout it, so let it lie
two days. Then take
Sage leaves, in fine powder 3 oz.
Garlic, minced finely 2 oz.
Shalots 2 oz.
Bay leaf, in powder 2 oz.
Chillies, or capsicum, in
powder 3 oz.
mix them well and then sift them, and blend them well with the meat,
so that all parts may partake of the flavour alike; put it into a jar,
which bung up close and set it aside for a week to get mellow. Now
try a small quantity of it: if it suits your taste, well and good, if not add
seasoning. Now fill your sausage skins, and when nice and solid
prick them a little, and put them into a pan of boiling water to simmer
slowly three quarters of an hour. Then take them out and let go cold,
and next day wipe them and smoke them three weeks with
Oak lops and dust 3 parts
Fern or grass turfs 3 parts
then hang them up in a dry room and keep them with hams, tongues,
&c. These will be excellent in a month, just popped into boiling water
for five minutes, or fried in olive oil and eaten cold.

jersey black puddings.

In France, in the Channel Islands, and Belgium these delicacies
are introduced at the tables of the highest families, a distinction
which they richly deserve, while in England very few persons make
them at home, but purchase at the shops an indescribable mass of
groats, blood, bread, herbs, &c., and frequently, to cover the
indolence of the pork-butcher, loaded with such an amount of the
commoner spices, as to render the whole anything but palatable.
Then, again, the fat, which seems to be the only recommendation, is
found here and there in lumps so large, that before they become
heated through, it is certain the other of part of the mass must be
burnt nearly to a cinder. I think that we can produce an article well
worthy of the trouble and slight expense incurred by the trial. Take a
couple of dozen of large onions, peel them, cutting off the bottoms,
cut them into small pieces, and put them into a stewpan along with
ten ounces of sweet lard, and stew them slowly till of a light brown
colour. Cut three pounds of pig’s leaf that is perfectly sweet and dry
into dice, pick out all the skins. Boil half a dozen heads of endive,
chop them fine and add to the fried onions; season them with
Table salt 2 oz.
 White pepper ¾ teaspoonful
Parsley, finely
chopped 4 tablespoonfuls
Thyme, in fine powder 1 tablespoonful
Bay leaves 2 tablespoonfuls
Half a nutmeg, grated
Now add three quarts of pig’s blood—calf’s, or sheep’s is as good—
that has not been long taken from the animal, and if still warm is
preferable. Mix all intimately, and if you find the mixture is too thin to
work well, add a handful or two of stale bread crumbs, or half to
three quarters of a pound of rice, boiled just tender but not mashed.
Take now the smaller pudding skins which have been properly
scraped and cleaned, wash them in salt and water, and with a tin
funnel tie one end of the skin tightly upon it, and the other end up in
a knot, two feet or a yard will be a convenient length. Proceed now to
fill your skins by pressing the meat through the funnel, pricking with a
pin to let out the air. When nicely and firmly filled, tie up the end, and
put the puddings into a flat large pan, with plenty of water tasting
slightly of salt, and already boiling, and let them simmer twenty
minutes, or rather more, attentively watching and pricking them to
prevent bursting. When the blood oozes out no longer, they are fully
cooked, then take them up and let them lie all night on a sieve. Cut
them into pieces four to six inches long, and when wanted for table
wipe them lightly over with a linen rag dipped in olive oil, and broil
them ten minutes. Serve them plain, but very hot. If you cannot
procure endive, you may substitute celery, which must be boiled
along with three or four laurel leaves; the tender parts only must be
used. Leeks also are generally liked, but in that case part of the
onions must be omitted.

marinated salmon.
Take six or eight pounds of the middle of a large fish, in
preference to a similar weight of the whole of a grilse or salmon
trout, scale it and cut off the fins, split it open at the back and reserve
the roe and liver; wipe the fish out nicely, and strew moist sugar all
over the red side, put it away on a dish until the next day. Then make
the following mixture,
Bay leaf, in powder 1 oz.
Mace, in powder ½ oz.
Cloves, in powder ¾ oz.
White pepper, in powder ¾ oz.
Table salt 1¼ lb.
Take out the backbone neatly and rub the fish on both sides well with
the mixture and the former sugar; lay three or four laurel leaves upon
the red face of the thick side, turn the thin side over upon them, and
set it away in a cool place for twenty-four hours. Then take it up,
wipe dry, and cut the sides into fitting pieces, and place them in a
deep dish upon the spices, and half a pint of Rhenish wine or Cape
Madeira; tie double paper over and bake it in a slow oven till done
enough. Pour off the pickle while yet warm, and set the pieces on a
sieve to drip until the next day—covered close with a cloth to prevent
the action of the air upon it, and thus to preserve the colour. Now
choose your pots according to the size of your family, and into each
one place a piece of the fish, the red face downwards, and fill up with
genuine olive oil, let remain until the next day, when add more oil,
and again on the third day replenishing what has been absorbed by
the fish. Then tie wetted bladder over the pots and keep them in a
cool dry air. This cannot fail of gaining the approval of all who
partake of it, and is the most certain method of obtaining fresh
salmon in the winter months. The same oil will do again and again
for a similar purpose.

marinated tench and carp.

How often have I seen twenty or thirty brace of these delicious
and exceedingly nutritious fish exposed for sale in some provincial
market-place, and spoiling for want of purchasers, some
neighbouring baronet or esquire having been drawing his pits and
preserves. But why no purchasers? Simply because, although the
spectators might nearly beg them, the price being so low, few
persons only would know what to do with them, they not being “poor
men’s fish;” and the sauces, &c., required to dress them would be
too expensive; the operatives have not seen pies made of them, like
eels, so even the million as they are termed do not enjoy them, and
ten to one the beautiful fish are lost; whilst if they were near
Houndsditch or the Minories, amongst the Israelites, they would not
remain half an hour unsold even at half-a-crown the brace, because
they know how to prepare them, and delicious indeed are the
majority of their fish dishes. Take three brace of these fish, scale
them, cut off the tails and fins, split them open at the back, take out
all the garbage, gills and eyes, and wash them quickly in salt and
water; now wipe them dry, and have ready the following mixture:
Fresh parsley, minced
finely 2 tablespoonfuls
Thyme, in powder 2 tablespoonfuls
Marjorum, in powder 2 tablespoonfuls
Moist sugar ½ lb.
Table salt ¾ lb.
Make two sides of each fish and cut them across, again place these
pieces in alternate layers with the mixture in an oblong pot closely
covered up, and set them aside for forty-eight hours; then unpack
the fish and replace them in the same receptacle, having rubbed
them well and turned each piece over; so let remain twenty-four
hours longer, the thickest pieces lying nearest the bottom of the jar.
Now unpack again and wash the pieces separately in weak vinegar
and water with a trifle of saltpetre in it, and wiping, leave them
between cloths while you are preparing, by boiling the used herbs
and liquor with half a pint of vinegar and water for ten minutes, and
skimming well, strain it through a fine sieve and set it aside. Now
take
Mace, bruised ½ oz.
Cloves, bruised 1 oz.
Cayenne pepper 1 teaspoonful
Twelve bay leaves, shred
 Four laurel leaves, green
Of the strained liquor ½ pint
Boil these for fifteen minutes, skim thoroughly, strain through a fine
sieve and add port wine half a pint, return this to the saucepan,
simmer it covered close for twelve minutes, and let remain by the
fireside. Lay your pieces of fish in a pie dish that will just hold them,
the thickest near the bottom, and pour the liquor in the saucepan
over the fish, tie paper doubled over the dish and bake till done
moderately, which can always be ascertained by the fish leaving the
bone upon a knife being applied. Now pour off the liquor, take out the
fish and set it to drain all night, covered up; place the pieces in the
best adapted pots and fill up with olive oil, renewing on the third day
what had been absorbed by the fish, and tie bladder over. In this and
similar cases take care that the oil is sufficient in quantity to cover
the contents to the thickness of three quarters of an inch at least.
These will be remarkably fine eating in a month. You cannot
purchase such at the shops.

marinated shrimps.
To ensure this delicacy of first-rate quality you must buy your
shrimps alive, and boil and pick them at home with all possible
despatch. Set a gallon of live shrimps in three quarts of boiling water,
in which one and half pounds of bay or rock salt has been dissolved,
and boil them fifteen minutes, then strain off the water and as soon
as possible commence shelling them, and keep them covered with
cloths as you proceed. Be particular that no refuse or pieces of the
shells remain to disgrace the preparer, and laying them lightly on
clean dishes, mix amongst them the following seasoning in very fine
powder, being well sifted and dried the day previous and kept bottled
up. I repeat that the shrimps must have been deprived of all their
moisture with the assistance of a moderately cool oven, and not two
fish to be seen sticking together. Nor can any excuse avail in this
instance, as the less time that is occupied in potting these sorts so
much better chance will there be of your total success in the product.
Mace, in finest powder 1 oz.
 Nutmeg, in finest powder 2 oz.
Cloves, in finest powder 2 oz.
Table salt, in finest powder 1 lb.
Loaf sugar, in finest
powder ¾ lb.
When seasoned to your entire satisfaction, the fish however fully
retaining their own fine flavour, fill your pots, which must be the
cleanest and best dried possible; press the shrimps well down, and
set them as you proceed at the mouth of your cool oven until there is
no evaporation observable. Now take them up, press down again,
and laying one or two bay leaves on the top, fill up with the best olive
oil, and set away in a cool dry room. Next day replenish the oil that
has been absorbed, and not till after the third day tie bladder over.
Plenty of oil must cover the fish, and although potted shrimps are not
expected to keep well after three or four days, yet you may expect
every satisfaction from this process. If decay soon takes place, it will
be owing to the fish not having been well dried in the process. As to
the preservation of the true flavour of the fish, this preparation
demands our decided preference.

marinated trout and grayling.

Fish about a pound weight each are the best for this purpose, but
provided that they are perfectly fresh, have a clear red gill, and bright
eye, we should be induced to thus preserve fish of any size, because
by this process the true flavour of the fish may be effectually
retained. Take two dozen fish, from twelve to twenty ounces each,
scale them, take off the heads, tails, and fins, open them at the belly,
take out the garbage, and washing them quickly in salt and water,
dry them with cloths and leave them covered up. Make the following
seasoning:
Table salt 1 lb.
Cayenne pepper ¾ oz.
Sifted loaf sugar ½ lb.
Dried parsley 1 oz.
 Thyme ½ oz.
All these must be in finest powder, and sifted, and with a good
mixture of them rub the insides and outsides of the fish well. If you
have any roes of them, wash them well, roll them in the powder and
place them inside the fish, bring the sides together, and tie round
with thread; place them in a convenient pot of earthenware, and with
half a pint of good sherry wine, paper tied closely over, bake them
sufficiently; then, while warm, pour off the gravy and let the fish lie by
till next day, basting them two or three times with the liquor, and
removing to the top those which laid on the bottom. Now pour off the
gravy, strain it, and put it aside bottled up close. Take out the fish
and pay the backbones inside with a camel’s-hair brush and this
mixture:
Essence of cassia 1 tablespoonful
Essence of mace 1 tablespoonful
Essence of nutmegs 2 tablespoonfuls
Essence of bays 3 tablespoonfuls
Lay the fish on their backs and repeat this brushing over the bone
two or three times, then lay the fish on their flat, mix the strained
liquor in the bottle with whatever you have remaining of your
essence mixture, and six tablespoonfuls of good old malt whisky,
and pour this over the fish; baste well with it two or three times daily
for a week or until they have imbibed a fine flavour of the ingredients,
and tying over with leather. Now dry the fish by wiping outside only,
and place pieces in fitting oblong pots and cover with olive oil of first
quality. (See Note, No. 11.) Replenish the oil, and after the third day
make safe with wetted bladder and remove to a cool dry room. In a
month they will be very good, but if you could keep some three
months they would be much improved.

marinated silver eels.

Take ten pounds of fine fresh river eels, each one and a half
pound to two pounds, skin them, cut off the heads, tails, and fins, lay
them open at the backs and remove all visible bones, and wash in
salt and water; then dry them, cut them into pieces four inches long,
and lay them in salt and water with a bit of saltpetre in it for ten
hours; then make a seasoning thus:
Allspice, ground finely 1 oz.
Cayenne pepper, ground
finely 1 teaspoonful
Bay leaf, in fine powder 1 oz.
Parsley, dried and rubbed
fine 2 oz.
Dry the fish well with cloths, rub them all over plentifully with this
mixture, and let them lie packed closely for twenty-four hours; roll
them up neat into little collars, tie them round, and bake them in a
wide-mouthed jar with a teacupful of water and vinegar at the
bottom. Next day pour off the liquor, take out the collars and put
them singly into white earthenware pots, and cover with olive oil,
observing to replenish the oil before finally closing up with bladder.

superior marinated silver eels, of very high flavour.

Get fresh eels of ten to fifteen ounces each, skin them, take off
heads, tails, and fins, open them at the bellies, and, clearing away all
refuse, wash them in salt and water and dry with cloths. Then for ten
pounds of fish take
Garlic, minced finely ½ oz.
Chillies, minced finely 1 oz.
Juniper berries, minced
finely ¾ oz.
Jamaica pepper, powdered 1 oz.
Bay salt 1 lb.
Saltpetre ¼ lb.
Vinegar 1 pint
Boil these twenty minutes, skimming well, and let go cold. Cut the
fish into pieces, place them in a deep dish and pour this pickle over
them, so let them lie till next day. Take out the fish and rub each
piece separately with the mixture, replace the fish in the vessel and
let them lie so for twenty-four hours longer. Now wipe them and hang
them up in a current of air for two days; then with a soft brush take
off the herbs and spice and roll up tightly each piece into a collar,
which secure with narrow tapes. Choose pots of white earthenware
that will each just hold a collar, and so as to allow of the fish being
totally surrounded by oil, and fill up with olive oil and replenishing as
before directed previous to finishing with wetted bladder. Keep these
three months at least, when they will be a fine relish at a moderate
expense. They should be boiled in hot water, and when cold enough
the tapes taken off and the fish eaten cold.

pickled smelts, preferred to anchovies.

This is an elegant preparation of the smaller fish, the large ones
being chosen for the dinner table. Cut open the fish at the belly, and
wipe them well out with damp cloths, lay in a light coat of the
following seasoning, the roes being replaced, and put them in layers
with the mixture alternately, in a deep jar wide at the top:
Saltpetre, in powder ½ oz.
Bay salt, in powder 9 oz.
Mace, in powder ½ oz.
Cloves, in powder 1 oz.
Black pepper, in
powder 1 oz.
Cochineal, in powder ¾ oz.
These must have been well mixed. Press the fish well down in the
jar, and boiling sufficient of the best pickling vinegar with bay leaves,
and four or five laurel leaves, pour it upon the fish when cold, and tie
leather over the jar. They will require three months at least to
become mellow, and are much improved by keeping twice that
length of time.
 pickled lobsters.
In our hot summer months, and when lobsters are plentiful, it
would be wise to save some that would come in opportunely for
sauce and many dishes, when lobsters are scarce and high priced.
Take fresh boiled lobsters, split them, take out the meat as whole as
you can, and make a seasoning of
Mace, in fine powder 1 oz.
Cayenne pepper ¾ oz.
Nutmeg, in fine powder 1 oz.
Table salt, in fine
powder 6 oz.
by well mixing them. Rub the meat well with this and equally, so that
no part is left undefended. Put the fish down in an earthen jar, and
repeat the rubbing for a day or two. Pack it then in small jars and
pour the following pickle over it, so that it may be covered to
thickness of an inch:
Best vinegar 1 pint
Chillies 1 oz.
Cucumber vinegar ½ pint
Sal prunelle ¾ oz.
and in similar proportions for each pint of vinegar used. Mind that the
vinegar pickle covers well, and then put over all as much olive oil as
will cover to the depth of half an inch. Tie wetted bladder over and
leather upon that for safe keeping. It will be well recommended after
a trial.

marinated herrings.
The freshest full-roed herrings must be taken for this purpose.
Scale twenty of the shortest and fattest you can get in the middle of
the season, take off the heads, tails, fins, and open them at the
bellies, clean all out, scrape the backbone, and washing the fish in
salt and water, wipe them dry and let them lie until next day, after
rubbing a dessert-spoonful of good moist sugar into the inside of
each fish. Lay them upon their backs in any vessel not too deep, and
cover the fish over with the following mixture—the roes should be
nicely washed, dried, and laid by the side of the fish:
Coarse sugar 1 lb.
Bay salt, in fine
powder ¾ lb.
Black pepper 1 oz.
Eschalots, minced 1 oz.
Nutmegs, grated 1 oz.
putting half the fish in one layer then a layer of the mixture, and then
a layer of fish above that, and the spices covering all; so let them lie
covered up with a cloth three days. Then wipe the fish dry, place the
roes inside, cut them across in two parts, tie up in rolls with calico,
and lay them with plenty of bay leaves and four laurel leaves in a
deep dish and bake them, just covered over with good sound porter;
then while warm pour off the liquor, and set the fish on a sieve to go
cold, and next day put each one into a white earthenware jar and
cover with olive oil with the same precautions as before. In two
months they will be mellow and rich and greatly liked. The porter in
which they were baked, being boiled twenty minutes with a dozen of
shalots and cayenne, will be a very good cold sauce for chops,
steaks, fish, &c.

marinated sprats.
These shall be superior to what were called “sardines in oil,”
some thirty years ago, and sold in London at 4s. 6d. at first in small
tin boxes, and afterwards at 2s. 6d., each tin containing about
sixteen sprats—for sprats they certainly were. Take a peck of fresh
sprats and pick out for your present use as many of the largest and
most sound as will suit your purpose. Pull off the heads, cut off the
fins and tails, and draw out the little guts, wash them quickly through
salt and water, and dry them between cloths. Have ready the
following:
 Mace, in finest powder 1 oz.
Cloves, in finest powder 2 oz.
Nutmeg, in finest
powder 1 oz.
Saltpetre, in finest
powder 3 oz.
Table salt, in finest
powder 1 lb.
Bay leaf, in finest
powder 3 oz.
with which rub each fish on the inside first, and then throughout the
bulk; put them into small pots of white earthenware, closely packed;
tie bladder over them, merely to keep out the water, and place them
in a stewpan, adding water to reach half way up the sides of the jars,
cover the pan and set it on a slow fire to simmer until the fish are
cooked; then take them up, remove the covers, and set them aside
till the next day, when, pouring off any oil or fat that may have been
given out in cooking, fill up with olive oil and tie bladders safely over.

potted smelts.
For this purpose the smaller fish will do very well. Cut them open
at the belly, clean out and wash in salt and water, having first cut off
the tails and fins; now season them inside with
Mace, in powder ½ oz.
Nutmeg, in powder 1 oz.
Table salt 3 oz.
Cayenne pepper ½ oz.
Bay leaf ¾ oz.
and let them lie till next day, when lay them in oblong pots and cover
them with clarified butter; bake them in a slow oven, and when done
enough pour off the butter, and drain effectually, and let them remain
twelve hours. Now warm up the butter again in a water bath, and
running it through a warmed sieve, pour the clear again over the fish,
adding more clarified butter to cover well. Tie bladder over and keep
dry.

potted lobsters.
Take a couple of fine lobsters, place them in a tub with plenty of
weak salt and water, and brush them well from froth and slime, then
wash in pure water and pop them into boiling water in which some
salt and saltpetre have been dissolved. When done lay them on a
sieve to get cold, and next day cut open the shells, crack the claws
and take out the meat, which then pick over carefully from skins and
specks, the coral also must be separated and examined. Now cut
the meat into dice and pound it finely in a mortar, till of a nice smooth
consistence, adding clarified butter at pleasure. Then season with
Mace, in fine powder ¾ oz.
Nutmeg, in fine powder 1 oz.
Bay leaf, in fine powder 1 oz.
White pepper, in fine
powder 1 oz.
Cloves, in fine powder ¾ oz.
and working the paste well proceed to fill white pots, and putting a
part of the coral into the middle of each pot. Press well in and set
aside for the night. Put the pots next into a slow oven, and let the
steam escape that might arise, and when you have got them as dry
as it is possible, without changing the colour of the preparation, set
them by to go cold; then cover with clarified butter effectually and
writing paper over the tops of the pots.

potted crabs.
To meet with well potted crabs is a very rare occurrence, and,
speaking generally, is attributable to the carelessness of the
preparer. Crabs are very often out of season, that is, cannot be got
without great trouble and expense, and they are very much liked;
these admissions furnish reasons why we should have some well
preserved. Take half a dozen eastern coast crabs—those of
Hartlepool and neighbourhood are always sound and well flavoured,
while those on the western coast and about Ireland are little worth—
plug up all the holes with wooden pegs, throw them into a tubful of
salt and water and brush them thoroughly, then wash in pure water
and put them into boiling water to be cooked; then take them up and
lay them to get cold; pull off the claws and put the large shells into a
slow oven after having run off any water that may have collected in
them, and when no more steam arises from the slow drying, set
them also to get cold. Take all the meat out of the claws, pick out
very carefully all the little skins and strings, cut it into dice and pound
it in a mortar with clarified butter until of a nice plastic mass, which
cover up in a cloth. Now attack the body shells, pick out all the solid
meat, and, setting the coral aside, throw out all the skins and refuse
you can find, and beat up this also with best Durham mustard, as for
the table, some cayenne, chillie vinegar, and table salt. Place some
of this at the bottom of each pot, and having ready this seasoning,
mix what is requisite with the meat from the claws:
Mace 1 oz.
Cloves 1½ oz.
Nutmeg 1 oz.
Bay leaf 1 oz.
Table salt 6 oz.
White pepper 2 oz.
Work this well into the mass, and having put a portion of the hard
coral into each pot, fill up with the seasoned claw meat. Set the pots
in a slow oven to evaporate what moisture you can, then set aside
and when cold pour clarified butter plentifully over, which, when cold,
must again be covered with writing paper. If your ingredients were
genuine, your butter sweet, and well managed, and the fish of prime
quality, you will have a choice or rare preparation of these generally
admired fish.

side of venison collared.

 As this sort of animal food can only be got at one season of the
year, I recommend any person who is fond of good eating at a
moderate cost, to get the two sides or flitches of a prime fat buck,
and cure them in the two different ways herein described, with full
assurance that the result will be perfectly satisfactory. Take a side
and hang it up in a free current of air as long as ever you can trust it,
wiping it daily with a coarse cloth, and dusting it over, particularly the
fat parts, with ground black pepper, to prevent the flies settling on it.
With a sharp knife take off the outer skin, and, removing all sinews,
bone it nicely; divide it next into pieces fit for collars, that is, so that
when rolled up the ends may be level and firm. Rub vinegar well
over the meat and leave it packed closely in a deep dish and
covered up till the next day. Make ready a mixture of
Mace, in fine powder 1 oz.
Cloves, in fine powder 2 oz.
Nutmeg, in fine powder 1 oz.
White pepper, in fine
powder 2 oz.
Bay salt ¾ lb.
Common or rock salt ½ lb.
Bay leaves, shred 3 oz.
and with this rub the meat, which should have been dried with cloths
well, and pack them down to lie forty-eight hours. Then take them
up, and cutting slices of hard fat bacon, half an inch wide and six or
eight inches long, insert them in channels cut in the venison and
about two and a half inches apart; this should be done with
consummate nicety. Now roll up the meat into collars and bind tightly
with broad tape, and sew strong unbleached calico all round the
length. Lay paper shavings in the bottom of a deep straight-sided jar,
and upon them six or eight laurel leaves and twice as many bay
leaves, and place the collars lightly upon them, then pouring in half a
pint of water, tie doubled paper over the jar and bake slowly for three
hours at least, or until nicely cooked; then while warm pour off the
gravy into a basin to get cold, and taking out the collars set them to
drain all night. Next day, after wiping them dry, place the rolls of meat
in a jar that will just hold them without squeezing, and pour clarified
mutton suet with sweet lard in the proportion of a quarter of a pound
of the latter to twelve ounces of the former, and when cold covering
the meat to the thickness of two inches. Tie wet bladder over and put
away in a cold airy room. This should not be broached under two
months, and then by placing the jar in a pan of boiling water for
twenty minutes you can take out a collar and put away the jar, taking
care that the meat is totally covered by the suet and made safe from
the air. Pop the collar into a saucepan of boiling water and salt for
fifteen minutes, then take it out, wipe it dry, and when nearly cold
take off the envelopes and set it by till next day. Serve it garnished
with fresh parsley and slices of lemon or pickled red beetroot, which
two latter may be eaten with it, and then I think you will confess that
this “common part of venison” is very first-rate eating.

the other side smoked.

Proceed as with the former side, and having rubbed it well with
vinegar, make a mixture of
Allspice, ground 3 oz.
Black pepper, ground 2 oz.
Eschalots, minced 2½ oz.
Garlic, minced 1½ oz.
Bay salt ¾ lb.
Coarse sugar ¾ lb.
Bay leaves 2 oz.
and rub all parts thoroughly, and let them lie placed down in a deep
pan for forty-eight hours; then take them up, wipe dry and lard them
well with shreds of the best sweet fat bacon and roll up tightly into
collars, which bind with tape and envelope in calico or thin canvas,
and stow them away in a deep jar that will hold them conveniently till
the next day. Boil up the used spices, herbs, and salts with as much
porter as you think will cover the meat, well skimming it till no more
scum arises, and pour it hot over the meat; so let it remain covered
with leather for a week. Then hang up the collars in a draught of air,
minding to turn them every morning regularly, or the neglect is
certain to be detected when brought to table. Next suspend them in
your chimney, and smoke them for a fortnight with
Oak lops or sawdust 2 parts
Beech chips 2 parts
Fern 1 part
Peat 1 part
When cold you can coat them with gelatine composition and keep
them in malt cooms. Slices cut off these collars and broiled will be
excellent in three months, or, if you choose to boil them and eat cold,
take care they are put on the fire in ready boiling water, as we do not
want venison broth. (See Note, No. 10.)

young pig collared.

Your porkman having supplied you with a short, round pig, say
about ten to twelve weeks old, the neck taken off close up to the
shoulders and split down the back, you will proceed to take out the
bones and gristles and wash it five minutes in salt and water, then
wipe dry and rub the following mixture—
Mace, beaten finely ½ oz.
Cloves, beaten finely 1 oz.
Nutmeg, beaten finely 1 oz.
Pepper, beaten finely 1 oz.
Salt, beaten finely 1½ lbs.
—in all parts well, and let them lie pressed closely for twenty-four
hours, then take them up, wipe dry, roll the one half up into a collar,
with tape and new linen, and boil it in a pickle made by boiling up the
former used spices—adding thereto
Six laurel leaves, green
Twelve bay leaves
Vinegar 1 pint
 Ginger, beaten 1 oz.
coarsely
Bay salt ¾ oz.
—until tender. Examine the collars when nearly cold, and tighten the
binding if requisite. Cut them through, in the middle, straight across,
put each roll into a jar just capable of holding it; boil up the pickle
again, adding vinegar and porter, if not enough to cover your meat
completely; tie bladder over and keep a month, when it will be very
rich, yet delicate. The pickle must have been well skimned and all fat
removed from it. The other half may be thus treated, making an
agreeable change, though this latter will be much higher flavoured:
Proceed as with the last, until having rolled it up in a nice tight collar,
you will make this pickle:
Sage leaves, shred ½ handful
Capsicums, mixed ½ oz.
Juniper berries,
bruised 1 oz.
Six laurel leaves, shred
Garlic, shred ½ oz.
Porter or ale 1 quart
Salt ½ lb.
Boil these twenty minutes, skim well and pour over the meat, placed
in a deep jar that will just hold it, and tying leather over let it remain
three weeks. Then take it up, wipe dry, hang it in a current of dry air
for a week, turning it daily, and smoke it with
Oak lops or sawdust 2 parts
Fern 2 parts
Beech or birch chips 2 parts
for three weeks. When cold you can coat it with gelatine, or if packed
with hams, tongues, &c. &c., in malt cooms, will be a high, much
esteemed article, after being kept a month or two to become mellow.
Care must be taken in the broiling, as the meat will be extremely rich
and tender.