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 Lincoln Taiz • Eduardo Zeiger
lan Max M0ller • Angus Murphy
 eo Contents

CHAPTER 1 The Endomembrane System 25

Plant and Cell Architecture 1 The endoplasmic reticulum is a network of
Plant life Processes: Unifying Principles
Plant Classification and life Cycles
Box 1.1 Evolutionary Relationships
among Plants 3
Plant life cycles altern ate between diploid
and haploid generation s 4
Overview of Plant Structure 6
Plant cells are surrou nded by rigid cell walls
Primary and secondary cell walls differ in
their components 7
The cellulose microfibrils and matrix polymers are
synthesized via different mechanisms 9
Plasmodesmata allow the free movement of
molecules between cells 10
New cells origin ate in dividing tissues
called meristems 11
Box 1.2 The Secondary Plant Body 14
Plant Cell Types 15
Dermal tissue covers the surfaces of plants
Ground tissue forms the bodies of plants 15
Vascular tissue forms transport networks between
different parts of the plant 17
Plant Cell Organelles 17
internal membranes 25
2 Vacuoles have diverse functions in plant cells 26
Oil bodies are lipid -storing organelles 27
2
Microbodies play specialized metabolic roles in leaves
and seeds 27
Independently Dividing Semiautonomous
Organelles 28
Proplastids mature into specialized plastids in
different plant tissues 31
6
Chloroplast and m itochondrial division are
independent of nuclear division 31
The Plant Cytoskeleton 32
The plant cytoskeleton consists of microtubules
and microfilaments 32
Actin, tubu lin, and their polymers are in constant flux
in the living cell 33
Microtubu le protofilaments first assemble into flat
sheets before curling into cylinders 34
Cytoskeletal motor proteins mediate cytoplasmic
streaming and directed organelle movement 34
15
Cell Cycle Regulation 37
Each phase of the cell cycle has a specific set of
biochemical and cellu lar activities 37
The cell cycle is regulated by cyclins and cyclin-
dependent kinases 38

Biological membranes are bilayers that Mitosis and cytokinesis involve both microtubules
contain proteins 18 and the endomembrane system 39

The Nucleus 20
Gene expression involves both transcription CHAPTER 2
and translation 23
Water and Plant Cells 45
Posttranslational regulation determines the
life span of proteins 23 Water in Plant life 46
viii Table of Contents

The Structure and Properties of Water 46 CHAPTER 3

Water is a polar molecule that forms Water Balance of Plants 65
hydrogen bonds 47
Water is an excellent solvent 47 Water in the Soil 66
Water has distinctive thermal properties relative A negative hydrostatic pressure in soil water lowers
to its size 48 soil water potential 66
Water molecules are h ighly cohesive 48 Water moves through the soil by bulk flow 67
Water has a high tensile strength 49
Water Absorption by Roots 68
Diffusion and Osmosis 51 Water moves in the root via the apoplast, symplast,
and transmembrane pathways 69
Diffusion is the net movement of molecules by
random thermal agitation 51 Solute accumulation in the xylem can generate
"root pressure" 70
Diffusion is most effective over short distances 52
Osmosis describes the net movement of water across a Water Transport through the Xylem 71
selectively permeable barrier 53 The xylem consists of two types of transport cells 71

Water Potential 53 Water moves through the xylem by pressure-driven

The chemical potential of water represents the
free-energy status of water 53
Th ree major factors contribute to cell water
potential 54
Water potentials can be measured 55
Water Potential of Plant Cells 55
Water enters the cell along a water potential
gradient 56
Water can also leave the cell in response to a water
potential gradient 56
Water potential and its components vary with growth
condition s and location within the plant 58
Cell Wall and Membrane Properties
Small ch anges in plant cell volume cause large
changes in turgor pressu re 58
The rate at w hich cells gain or lose water is influenced
by plasma membrane hydraulic conductivity 60
Aquaporins facilitate the movement of water across
plasma membranes 60
Plant Water Status 61
Physiological processes are affected by plant
water status 61
Solute accumulation helps cells maintain turgor
and volume 62
bulk flow 73
Water movement th rough the xylem requires a
smaller pressure gradient than movement
through living cells 73
W hat pressure difference is needed to lift water 100
meters to a treetop? 74
The cohesion-ten sion theory explains water transport
in the xylem 74
Xylem transport of water in trees faces physical
challenges 76
Plants minimize the consequences of xylem
cavitation 78
Water Movement from the Leaf
58 to the Atmosphere 78
Leaves have a large hydraulic resistance 79
The driving force for transpiration is the difference
in water vapor concentration 80
Water loss is also regulated by the pathway
resistances 81
The bou ndary layer contributes to diffusional
resistance 81
Stomatal resistance is another major component
of diffusion al resistance 82
The cell walls of guard cells have specialized
features 82
An increase in guard cell turgor pressu re opens
the stomata 83
 •
Table of Contents IX

Coupling Leaf Transpiration and CHAPTER 5

Photosynthesis: Light-dependent Assimilation of Inorganic
Stomatal Opening 85 Nutrients 121
Stomatal opening is regulated by light 85
Stomatal opening is specifically regulated by Nitrogen in the Environment 122
blue light 86 Nitrogen passes through several forms in a
biogeochemical cycle 122
Water-use efficiency 87
Unassimilated ammonium or nitrate may be
dangerous 124
Overview: The Soil-Plant-Atmosphere
Continuum 87 Nitrate Assimilation 125
Many factors regulate nitrate reductase 125
Nitrite reductase converts nitrite to ammonium 126
CHAPTER 4
Both roots and shoots assimilate nitrate 127
Mineral Nutrition 91
Ammonium Assimilation 128
Essential Nutrients, Deficiencies,
Converting ammoniu m to amino acids requires
and Plant Disorders 92
two enzymes 128
Special tech niques are used in nutritional studies 95
Ammonium can be assimilated via an alternative
Nutrient solutions can sustain rapid plant growth 95 pathway 128
Mineral deficiencies disrupt plant metabolism and Transamination reactions transfer nitrogen 129
function 98
Asparagine and glutamine lin k carbon and
Analysis of plant tissues reveals mineral nitrogen metabolism 130
deficiencies 103
Amino Acid Biosynthesis 130
Treating Nutritional Deficiencies 104
Crop yields can be improved by the addition Biological Nitrogen Fixation 131
of fertilizers 105 Free-living and symbiotic bacteria fix nitrogen 131
Some mineral nutrients can be absorbed by leaves 106 Nitrogen fixation requires microanaerobic or
anaerobic conditions 132
Soil, Roots, and Microbes 106
Symbiotic nitrogen fixation occurs in specialized
Negatively charged soil particles affect the adsorption structures 134
of mineral nutrients 107
Establishing symbiosis requires an exchange
Soil pH affects nutrient availability, soil microbes, of signals 134
and root growth 108
Nod factors produced by bacteria act as signals
Excess mineral ions in the soil limit plant growth 109 for symbiosis 135
Some plants develop extensive root systems 109 Nodule formation involves phytohormones 136
Root systems differ in form but are based on The nitrogenase en zyme complex fixes N 2 138
common structures 110
Amides and ureides are the transported forms
Different areas of the root absorb different of n itrogen 139
mineral ions 112
Nutrient availability influences root growth 114 Sulfur Assimilation 140
Mycorrhizal symbioses facilitate nutrient uptake Sulfate is the form of sulfur transported
by roots 114 into plants 140
Nutrients move between mycorrhizal fungi and Sulfate assimilation occurs mostly in leaves 140
root cells 118 Methionine is synthesized from cysteine 141
x Table of Contents

Phosphate Assimilation 141 Plasma membrane H+-ATPases are highly regulated

P-type ATPases 170
Iron Assimilation 141 The tonoplast H+-ATPase drives solute accumulation
Roots modify the rhizosphere to acquire iron 141 in vacuoles 171
Iron cations form complexes with carbon H+-pyrophosphatases also pump protons at
and phosphate 142 the tonoplast 173

The Energetics of Nutrient lon Transport in Stomatal Opening 173

Assimilation 143 Light stimulates ATPase activity and creates a
stronger electrochemical gradient across the guard
cell plasma membrane 173
CHAPTER 6 Hyperpolarization of the guard cell plasma membrane
leads to uptake of ions and water 175
Solute Transport 147
lon Transport in Roots 176
Passive and Active Transport 148
Solutes move through both apoplast and symplast 176
Transport of Ions across Ions cross both symplast and apoplast 176
Membrane Barriers 150 Xylem parenchyma cells participate in
Different diffusion rates for cations and anions xylem loading 178
produce diffusion potentials 151
How does membrane potential relate to ion
distribution? 152 CHAPTER 7
The N ernst equation distinguishes between active Photosynthesis: The Light
and passive transport 153
Reactions 181
Proton transport is a major determinant of the
membrane potential 154
Photosynthesis in Higher Plants 182
Membrane Transport Processes 155 General Concepts 182
Ch annels en hance diffusion across membranes 156 Light has characteristics of both a particle and
Carriers bind and transport specific substances 158 a wave 182
Primary active transport requires energy 158 When molecules absorb or emit light, they change
their electronic state 183
Secondary active transport uses stored energy 160
Photosynthetic pigments absorb the light th at
Kinetic analyses can elucidate transport
powers photosynthesis 185
mechanisms 161

Membrane Transport Proteins 163 Key Experiments in Understanding

Photosynthesis 186
The genes for many transporters have been
identified 163 Action spectra relate light absorption to
photosynthetic activity 186
Transporters exist for diverse n itrogen-containing
compounds 165 Photosynthesis takes place in complexes containing
light-h arvesting antennas and photochemical
Cation transporters are diverse 166 reaction centers 187
Anion transporters have been identified 168 The chemical reaction of photosynthesis is driven
Transporters for metal and metalloid ions transport by light 189
essential micronutrients 169 Light drives the reduction of NADp+ and the
Aquaporins have diverse functions 170 formation of ATP 189
Oxygen-evolving organisms have two photosystems
that operate in series 190

Organization of the
Photosynthetic Apparatus 192
The chloroplast is the site of photosynthesis 192
Thylakoids contain integral membrane
proteins 192
Photosystems I and II are spatially separated in
the thylakoid membrane 193
Organization of Light-Absorbing
Antenna Systems 195
Antenna systems contain chlorophyll and are
membrane-associated 195
The antenna funnels energy to the
reaction center 195
Many antenna pigment- protein complexes have
a common structural motif 196
Mechanisms of Electron
Transport 197
Electrons from chlorophyll travel through
the carriers organized in the Z scheme 197
Energy is captured when an excited chlorophyll
reduces an electron acceptor molecule 199
The reaction center chlorophylls of the
two photosystems absorb at different
wavelengths 200
The PSII reaction center is a multi-subunit
pigment- protein complex 201
Water is oxidized to oxygen by PSII 202
Pheophytin and two quinones accept electrons
from PSII 202
Electron flow through the cytochrome b6f complex
also transports protons 203
Plastoquinone and plastocyanin carry
electrons between photosystem II and
photosystem I 205
The PSI reaction center reduces NADpt 205
Cyclic electron flow generates ATP but no
NADPH 206
Some herbicides block photosynthetic electron
flow 206
Proton Transport and ATP Synthesis
in the Chloroplast 207
•
Table of Contents XI
CHAPTER 8
Photosynthesis: The Carbon
Reactions 213
The Calvin-Benson Cycle 214
The Calvin- Benson cycle has three phases:
carboxylation, reduction, and regeneration 215
The fixation of C02 via carboxylation of
ribulose 1,5-bisphosphate and the reduction
of the product 3-phosphoglycerate yield
triose phosphates 215
The regeneration of ribulose 1,5-bisphosphate
ensures the continuous assimilation of C02 217
An induction period precedes the steady state of
photosynthetic C02 assimilation 219
Many mechanisms regulate the Calvin- Benson
cycle 220
Rubisco activase regulates the catalytic activity
of Rubisco 221
Light regulates the Calvin- Benson cycle via the
ferredoxin- thioredoxin system 221
Light-dependent ion movements modulate enzymes
of the Calvin- Benson cycle 222
Photorespiration: The C2 Oxidative
Photosynthetic Carbon Cycle 222
The oxygenation of ribulose 1,5-bisphosphate
sets in motion the c2 oxidative photosynthetic
carbon cycle 224
Photorespiration is linked to the photosynthetic
electron transport chain 227
Inorganic Carbon-Concentrating
Mechanisms 228
Inorganic Carbon-Concentrating
Mechanisms: The C4 Carbon Cycle 229
Malate and aspartate are the primary
carboxylation products of the c4 cycle 229
The C4 cycle assimilates C02 by the concerted
action of two different types of cells 229
Bundle sheath cells and mesophyll cells exhibit
anatomical and biochemical differences 231
The C4 cycle also concentrates C02 in
single cells 232
xii Table of Contents
Light regulates the activity of key C4
enzymes 234
Photosynthetic assimilation of C0 2 in C4
plants demands more transport processes
than in c3plants 234
In hot, dry climates, the C4 cycle reduces
photorespiration 234
Inorganic Carbon-Concentrating
Mechanisms: Crassulacean Acid
Metabolism (CAM) 235
Different mechanisms regulate C4 PEPCase
and CAM PEPCase 237
CAM is a versatile mechan ism sensitive to
environmental stimuli 23 7
Accumulation and Partitioning
of Photosynthates-Starch and
Sucrose 238
CHAPTER 9
Photosynthesis: Physiological and
Ecological Considerations 243
The Effect of Leaf Properties on
Photosynthesis 245
Leaf anatomy and canopy structu re maximize
light absorption 245
Leaf angle and leaf movement can control
light absorption 248
Leaves acclimate to sun and shade environments
Effects of Light on Photosynthesis
in the Intact Leaf 250
Light-response curves reveal photosynthetic
properties 250
Leaves must dissipate excess light energy 252
Absorption of too much light can lead to
photoinhibition 255
Effects of Temperature on Photosynthesis
in the Intact Leaf 256
Leaves must dissipate vast quantities of heat 256
There is an optimal temperature for
photosynthesis 257
Photosynthesis is sensitive to both high and low
temperatures 257
Photosynthetic efficiency is
temperatu re-sen sitive 258
Effects of Carbon Dioxide on
Photosynthesis in the Intact Leaf 259
Atmospheric C0 2 concentration keeps rising 259
C02 diffusion to the chloroplast is essential to
photosynthesis 260
C02 imposes limitations on photosynthesis 261
H ow will photosynthesis and respiration change in
the future u nder elevated C02 condition s? 264
CHAPTER 10
Translocation in the Phloem 269
Patterns of Translocation:
Source to Sink 270
Pathways of Translocation 271
Sugar is translocated in phloem sieve elements 272
Mature sieve elements are living cells specialized
for translocation 272
Large pores in cell walls are the prominent featu re
of sieve elements 274
Damaged sieve elements are sealed off 275
Companion cells aid the h ighly specialized sieve
elements 276
249
Materials Translocated in the Phloem 276
Phloem sap can be collected and analyzed 277
Sugars are translocated in a nonreducing form 278
O ther solutes are tran slocated in the phloem 278
Rates of Movement 280
The Pressure-Flow Model, a Passive
Mechanism for Phloem Transport 280
An osmotically generated pressure gradient drives
tran slocation in the pressu re-flow model 281
Some predictions of pressure flow h ave been
confirmed, while others require further
experimentation 282
 •••
Table of Contents XIII

There is no bidirectional transport in single sieve Transport of Signaling Molecules 298

elements, and solutes and water move at the
Turgor pressure and chemical signals coordinate
same velocity 283
source and sink activities 298
The energy requirement for transport through the
Proteins and RNAs function as signal molecules
phloem pathway is small in herbaceous plants 283
in the phloem to regulate growth and
Sieve plate pores appear to be open channels 284 development 299
Pressure gradients in the sieve elements may be Plasmodesmata function in phloem signaling 299
modest; pressures in herbaceous plants and
trees appear to be similar 284

Phloem Loading 285 CHAPTER 11

Phloem loading can occur via the apoplast Respiration and Lipid
or symplast 285 Metabolism 303
Abundant data support the existence of apoplastic
loading in some species 287 Overview of Plant Respiration 303
Sucrose uptake in the apoplastic pathway requires
metabolic energy 287
Glycolysis 306
Glycolysis metabolizes carbohydrates from
Phloem loading in the apoplastic pathway involves a
several sources 307
sucrose- H+ symporter 288
The energy-conserving phase of glycolysis extracts
Phloem loading is symplastic in some species 288
usable energy 309
The polymer-trapping model explains symplastic
Plants have alternative glycolytic reactions 310
loading in plants with intermediary-type
companion cells 288 In the absence of oxygen, fermentation regenerates
the NAD+ needed for glycolytic ATP
Phloem loading is passive in several tree species 290
production 310
Phloem Unloading and
The Oxidative Pentose Phosphate
Sink-to-Source Transition 290
Pathway 311
Phloem unloading and short-distance
The oxidative pentose phosphate pathway produces
transport can occur via symplastic or
NADPH and biosynthetic intermediates 313
apoplastic pathways 290
The oxidative pentose phosphate pathway is
Transport into sink tissues requires
redox-regulated 313
metabolic energy 291
The transition of a leaf from sink to source The Tricarboxylic Acid Cycle 314
is gradual 292
Mitochondria are semiautonomous organelles 314
Photosynthate Distribution: Pyruvate enters the mitochondrion and is oxidized
Allocation and Partitioning 294 via the TCA cycle 315

Allocation includes storage, utilization, The TCA cycle of plants has unique features 317
and transport 294
Mitochondrial Electron Transport
Various sinks partition transport sugars 295
and ATP Synthesis 318
Source leaves regulate allocation 295
The electron transport chain catalyzes a flow of
Sink tissues compete for available translocated electrons from NADH to 0 2 318
photosynthate 297
The electron transport chain has supplementary
Sink strength depends on sink size branches 320
and activity 297
ATP synthesis in the mitochondrion is coupled to
The source adjusts over the long term to changes electron transport 321
in the source-to-sink ratio 298
xiv Table of Contents
Tran sporters exchange substrates and products 322
Aerobic respiration yields about 60 molecules
of ATP per molecule of sucrose 324
Plants h ave several mech anisms that lower the ATP
yield 324
Short-term control of mitochondrial respiration
occurs at different levels 326
Respiration is tightly coupled to other pathways 328
Respiration in Intact Plants and Tissues 329
Plants respire roughly half of the daily
photosynthetic yield 329
Respiratory processes operate during
photosynthesis 329
Different tissues and organs respire at
different rates 330
Environ mental factors alter respiration rates 331
Lipid Metabolism 332
Fats and oils store large amounts of energy 333
Triacylglycerols are stored in oil bodies 333
Polar glycerolipids are the main structural lipids
in membranes 334
Membrane lipids are precursors of important
sign aling compounds 334
Storage lipids are converted into carbohydrates in
germinating seeds, releasin g stored energy 336
CHAPTER 12
Signals and Signal Transduction 341
Temporal and Spatial Aspects
of Signaling 342
Signal Perception and Amplification 343
Signals must be amplified intracellularly to regulate
their target molecules 344
Ca2+ is the most ubiquitous second messenger in
plants and other eukaryotes 344
Ch anges in the cytosolic or cell wall pH can serve
as second messengers for hormonal and stress
responses 345
Reactive oxygen species act as second messengers
mediating both environmental and developmental
sign als 347
Hormones and Plant Development 347
Auxin was discovered in early studies of coleoptile
bending du ring phototropism 349
Gibberellins promote stem growth and were
discovered in relation to the "foolish seedling
disease" of rice 349
Cytokin ins were discovered as cell division-
promoting factors in tissue culture
experiments 351
Ethylene is a gaseous hormone th at promotes fruit
ripening and other developmental processes 351
Abscisic acid regulates seed maturation and stomatal
closure in response to water stress 351
Brassinosteroids regulate floral sex determination,
photomorphogenesis, and germin ation 352
Salicylic acid and jasmonates function in defense
responses 353
Strigolactones suppress branching and promote
rh izosphere interactions 353
Phytohormone Metabolism
and Homeostasis 353
Indole-3-pyruvate is the primary intermediate in
auxin biosynthesis 354
Gibberellins are synthesized by oxidation of the
diterpene ent-kaurene 354
Cytokin ins are adenine derivatives w ith isoprene
side chains 355
Ethylene is synthesized from meth ionine via the
intermediate ACC 355
Abscisic acid is synthesized from a carotenoid
intermediate 355
Brassinosteroids are derived from the sterol
campesterol 356
Strigolactones are synthesized from
~ - carotene 358
Signal Transmission and Cell-Cell
Communication 358
Hormonal Signaling Pathways 359
The cytokinin and ethylene signal transduction
pathways are derived from the bacterial two-
component regulatory system 359
Receptor-like kin ases mediate brassinosteroid
signaling 360
 Table of Contents xv

The core ABA sign aling components include
phosphatases and kinases 362
Plant hormone signaling pathways generally employ
negative regulation 362
Protein degradation via ubiquitin ation plays a
prominent role in hormone signaling 362
Plants h ave mech anisms for switching off or
attenu ating signaling respon ses 363
The cellular response output to a signal is often
tissue-specific 363
Cross-regulation allows signal tran sduction pathways
to be integrated 363
The nucleus is a primary site of cryptochrome
action 382
Cryptochrome interacts with phytochrome 382
Phototropins 383
Phototropism requires changes in auxin
mobilization 384
Phototropin s regulate chloroplast movements 384
Stomatal opening is regulated by blue light which
activates the plasma membrane H+ -ATPase 385
The Coaction of Phytochrome,
Cryptochrome, and Phototropins 386

Responses to Ultraviolet Radiation 387

CHAPTER 13
Signals from Sunlight 369
CHAPTER 14
Plant Photoreceptors 372
Embryogenesis 391
Photoresponses are driven by light quality or spectral
properties of the energy absorbed 372 Overview of Embryogenesis 393
Plant responses to light can be distinguished by the
amount of light required 375 Comparative Embryology of Eudicots and
Monocots 393
Phytochromes 375 Morphological similarities and differences between
Phytoch rome is the primary photoreceptor for red eudicot and monocot embryos dictate their
and far-red light 375 respective patterns of development 394
Phytoch rome can interconvert between Pr and Pfr Apical-basal polarity is maintained in the embryo
forms 376 du ring organogenesis 396
Embryo development requires regulated
Phytochrome Responses 377 communication between cells 398
Phytoch rome responses vary in lag time and escape
Auxin sign aling is essential for embryo
time 377
development 400
Phytoch rome responses fall into th ree main categories
Polar auxin tran sport is mediated by localized auxin
based on the amount of light required 378
efflu x carriers 401
Phytoch rome A mediates respon ses to continuous
Auxin synthesis and polar transport regulate
far-red light 379
embryon ic development 404
Phytoch rome regulates gene expression 380
Radial patterning guides formation of tissue
layers 404
Blue-Light Responses and
Photoreceptors 380 The protoderm differentiates into the epidermis 405

Blue-light responses have ch aracteristic kinetics and The central vascular cylinder is elaborated by
lag times 381 cytokin in-regulated progressive cell divisions 405

Cryptochromes 381 Formation and Maintenance

Blue-light irradiation of the cryptochrome FAD
of Apical Meristems 406
chromophore causes a con formational ch ange 382 Auxin and cytokinin contribute to the formation
and maintenance of the RAM 406
xvi Table of Contents

SAM formation is also influenced by factors involved Vascular differentiation begins during
in auxin movement and responses 407 seedling emergence 427
Cell proliferation in the SAM is regulated by cytokinin Growing roots h ave distinct zones 427
and gibberellin 408 Ethylene and other hormones regulate
root h air development 428
Lateral roots arise internally from the pericycle 428
CHAPTER 15
Seed Dormancy, Germination, Cell Expansion: Mechanisms
and Hormonal Controls 430
and Seedling Establishment 411
The rigid primary cell wall must be loosened for cell
Seed Structure 412 expansion to occu r 430
Seed anatomy varies widely among different plant Microfibril orientation influences growth
groups 412 direction ality of cells with diffuse growth 431
Acid-induced growth and cell wall yielding are
Seed Dormancy 415 mediated by expansin s 432
There are two basic types of seed dormancy Auxin promotes growth in stems and coleoptiles,
mechanisms: exogenous and endogenous 415 while in hibiting growth in roots 433
Non-dormant seeds can exhibit vivipary The outer tissues of eudicot stems are the targets of
and precocious germination 416 auxin action 433
The ABA:GA ratio is the primary determin ant The minimum lag time for auxin-induced elongation
of seed dormancy 417 is 10 minutes 434
Auxin -induced proton extrusion loosens
Release from Dormancy 419
the cell wall 434
Light is an important signal th at breaks dormancy in
small seeds 419 Ethylene affects microtubule orientation and induces
lateral cell expansion 435
Some seeds require either chilling or after-ripening
to break dormancy 419 Tropisms: Growth in Response
Seed dormancy can be broken by various chemical to Directional Stimuli 435
compou nds 420 Auxin transport is polar and gravity-independent 436

Seed Germination 421 The Cholodny-Went hypothesis is supported by

Germination and postgermin ation can be divided
into three phases corresponding to the phases
of water uptake 421
Mobilization of Stored Reserves 423
The cereal aleurone layer is a specialized digestive
tissue surrounding the starchy endosperm 423
Seedling Establishment 425
The development of emerging seedlings is strongly
influenced by light 425
auxin movements and au xin responses during
gravitropic growth 437
Gravity perception is triggered by the sedimentation
of amyloplasts 438
Gravity sensing may involve pH and calcium ions
(Ca2+) as second messengers 439
Phototropin s are the light receptors involved in
phototropism 440
Phototropism is mediated by the lateral redistribution
of auxin 440

Gibberellins and brassinosteroids both suppress Shoot phototropism occurs in a series of steps 441
photomorphogenesis in darkness 426
Hook opening is regulated by phytoch rome, auxin,
and ethylene 426
 ••
Table of Contents XVII

CHAPTER 16 Leaf senescence may be sequential, seasonal,

or stress-induced 463
Vegetative Growth and
The earliest cellular changes du ring leaf senescence
Senescence 445 occur in the chloroplast 464

The Shoot Apical Meristem 445 Reactive oxygen species serve as internal signaling
agents in leaf senescence 464
The shoot apical meristem has distinct zones and
layers 446 Plant hormones interact in the regulation
ofleafsenescence 465
leaf Structure and Phyllotaxy 447
leaf Abscission 467
Auxin-dependent patterning of the shoot apex begins
during embryogenesis 448 The timing of leaf abscission is regulated by the
interaction of ethylene and auxin 46 7
Differentiation of
Epidermal Cell Types 450 Whole Plant Senescence 469
A specialized epidermal lineage produces guard Angiosperm life cycles may be annual, biennial,
cells 451 or perennial 469
Nutrient or hormonal redistribution may trigger
Venation Patterns in leaves 452 senescence in monocarpic plants 470
The primary leaf vein is initiated in the leaf
primordium 452
Auxin canalization initiates development of the leaf CHAPTER 17
trace 452
Flowering and Fruit
Shoot Branching and Architecture 454 Development 473
Auxin, cytokinins, and strigolactones regulate axillary
bud outgrowth 454
Floral Evocation: Integrating Environmental
Cues 474
The initial signal for axillary bud growth may be an
increase in sucrose availability to the bud 456 The Shoot Apex and Phase Changes 474
Shade Avoidance 457 Plant development has th ree phases 475

Reducing shade avoidance responses can improve
crop yields 458
Root System Architecture 458
Plants can modify their root system architecture
to optimize water and nutrient uptake 458
Monocots and eudicots differ in their
root system architectu re 458
Root system architectu re changes in response to
phosphorus deficiencies 459
Plant Senescence 461
During leaf senescence, nutrients are remobilized
from the source leaf to vegetative or reproductive
sinks 462
Juvenile tissues are produced first and are located at
the base of the shoot 475
Phase changes can be influenced by nutrients,
gibberellins, and epigenetic regulation 476
Photoperiodism: Monitoring
Day length 476
Plants can be classified according to their
photoperiodic responses 477
Photoperiodism is one of many plant processes
controlled by a circadian rhythm 479
Circadian rhythms exhibit characteristic features 479
Circadian rhythms adjust to different
day-night cycles 481

The developmental age of a leaf may differ from its The leaf is the site of perception of the
chronological age 462 photoperiodic signal 482
 •••
XVIII Table of Contents

Plants monitor day length by measuring the length Fruit Development and Ripening 498
of the night 482
Arabidopsis and tomato are model systems for the
Night breaks can cancel the effect of the study of fr uit development 498
dark period 482
Fleshy fr uits undergo ripening 500
Photoperiodic timekeeping during the n ight depends
Ripen ing involves changes in the color of fruit 500
on a circadian clock 483
Fruit softening involves the coordinated action of
A coincidence model links oscillating light sensitivity
many cell wall- degrading enzymes 501
and photoperiodism 484
Taste and flavor reflect changes in acids, sugars,
Phytochrome is the primary photoreceptor in
and aroma compounds 502
photoperiodism 486
The causal link between ethylene and ripening
Vernalization: Promoting was demonstrated in transgenic and mutant
Flowering with Cold 487 tomatoes 502
Climacteric and non-climacteric fr uits differ in their
Long-distance Signaling ethylene responses 503
Involved in Flowering 488
Gibberellins and ethylene can induce
flowering 489 CHAPTER 18
Floral Meristems and Floral Biotic Interactions 507
Organ Development 490
Beneficial Interactions between Plants
The SAM in Arabidopsis changes with
development 491
and Microorganisms 509
Other types of rhizobacteria can increase nutrient
The fo ur different types of floral organs are initiated
availability, stimulate root branching, and protect
as separate whorls 491
against pathogens 509
Two major categories of genes regulate floral
development 492 Harmful Interactions of Pathogens
The ABC model partially explains the determination and Herbivores with Plants 510
of floral organ identity 492 Mech anical barriers provide a first line of defense
against insect pests and pathogens 511
Pollen Development 494
Specialized plant metabolites can deter insect
herbivores and pathogen in fection 513
Female Gametophyte Development
in the Ovule 495 Plants store constitutive toxic compounds in
specialized structures 514
Functional megaspores undergo a series
of free nuclear mitotic division s followed Plants often store defensive chemicals as nontoxic
by cellularization 495 water-soluble sugar conjugates in specialized
vacuoles 517
Pollination and Double Fertilization
in Flowering Plants 496 Inducible Defense Responses
Two sperm cells are delivered to the female
to Insect Herbivores 519
gametophyte by the pollen t ube 497 Plants can recognize specific components of
insect saliva 519
Pollination begins with adhesion and hydration of a
pollen grain on a compatible flower 497 Ph loem feeders activate defense signaling
pathways similar to those activated by
Pollen tubes grow by tip growth 497
pathogen infections 520
Double fertilization results in the formation of the
zygote and the primary endosperm cell 498

Jasmonic acid activates defense responses against
insect herbivores 520
Hormonal interactions contribute to plant-insect
herbivore interactions 521
JA initiates the production of defense proteins
that in hibit herbivore digestion 521
Herbivore damage induces systemic defenses 522
Long-distance electrical signaling occurs in response
to insect herbivory 522
Herbivore-induced volatiles can repel herbivores
and attract natural enemies 524
Herbivore-induced volatiles can serve as long-distance
signals within and between plants 525
Insects have evolved mechanisms to defeat
plant defenses 526
Plant Defenses against Pathogens 526
Microbial pathogens have evolved various strategies
to invade host plants 526
Pathogens produce effector molecules that aid in the
colonization of their plant host cells 527
Pathogen infection can give rise to molecular "danger
signals" that are perceived by cell surface pattern
recognition receptors (PRRs) 528
R proteins provide resistance to individual pathogens
by recognizing strain -specific effectors 528
The hypersensitive response is a common defense
against pathogens 530
A single encounter with a pathogen may increase
resistance to future attacks 531
Plant Defenses against
Other Organisms 531
Some plant parasitic nematodes form specific
associations through the formation of distinct
feeding structu res 531
Plants compete with other plants by secreting
allelopathic secondary metabolites into the
soil 533
Some plants are biotrophic pathogens of
other plants 533
•
Table of Contents XIX
CHAPTER 19
Abiotic Stress 537
Defining Plant Stress 538
Physiological adjustment to abiotic stress involves
trade-offs between vegetative and reproductive
development 539
Acclimation versus Adaptation 540
Environmental Stressors 541
Water deficit decreases turgor pressure, increases
ion toxicity, and in hibits photosynthesis 541
Salinity stress has both osmotic and cytotoxic
effects 543
Temperature stress affects a broad spectru m
of physiological processes 543
Flooding results in anaerobic stress to the root 544
Light stress can occur when shade-adapted or
shade-acclimated plants are subjected to full
su nlight 544
Heavy metal ions can both mimic essential mineral
nutrients and generate ROS 545
Combinations of abiotic stresses can induce unique
signaling and metabolic pathways 545
Sequential exposure to different abiotic stresses
sometimes confers cross-protection 547
Plants use a variety of mechanisms to sense
abiotic stress 547
Physiological Mechanisms That Protect
Plants against Abiotic Stress 548
Plants can alter their morphology in response to
abiotic stress 548
Metabolic shifts enable plants to cope with a variety of
abiotic stresses 549
Heat shock proteins maintain protein integrity under
stress conditions 549
Membrane lipid composition can adjust to changes in
temperature and other abiotic stresses 550
Chloroplast genes respond to high-intensity light by
sending stress signals to the nucleus 551
A self-propagating wave of ROS mediates systemic
acquired acclimation 551
Abscisic acid and cytokinins are stress-response
hormones that regulate drought responses 552
xx Table of Contents

Plants adjust osmotically to drying soil by
accumulating solutes 553
Epigenetic mechanisms and small RNAs provide
additional protection against stress 555
Submerged organs develop aerenchyma tissue in
response to hypoxia 556
Antioxidants and ROS -scavenging pathways protect
cells from oxidative stress 558
Exclusion and internal tolerance mechanisms
allow plants to cope w ith toxic metal and
metalloid ions 559
Plants use cryoprotectant molecules and antifreeze
proteins to prevent ice crystal formation 560

Glossary G-1
Illustration Credits IC-1
Index 1-1
Pre ace
Over the course of publishing six editions of Plant Physiol-
ogy (now Plant Physiology and Development), it has become
abundantly clear to us that, as a consequence of the spec-
tacular advances in plant developmental genetics, the field
of plant physiology has broadened in scope and increased
in depth to the p oint where it is no longer possible for a
sin gle textbook to serve the needs of all cou rses on plant
physiology.
Based on extensive feedback from instructors affiliated
with a wide range of u niversities and colleges, we designed
Fundamentals of Plant Physiology for in structors seeking a
shorter text that emphasizes concepts as opposed to detailed,
comprehensive coverage. To accommodate instructors
wishing to focu s their lectures on canonical plant physiol-
ogy topics, the treatment of plant development, including
hormonal sign aling and photoreceptors, has been greatly
condensed and reorganized. In keeping with a conceptual
approach, genetic pathways have been streamlin ed by
prioritizing functional mechan isms and by substituting
descrip tive ter ms for three-letter gene n ames whenever
possible.
Th roughout the text, explanation s have been revised
for easier comprehension, and figures have been simplified
and annotated for greater clarity. To further improve read-
ability, we have switched to a single-column format that
includes a running glossary in the margin s. In working
on Fundamentals of Plant Physiology, our primary goal has
been to provide a more concise, accessible, and focused
treatment of the field of plant physiology, while at the same
time maintaining the high standard of scientific accuracy
and pedagogical richness for which Plant Physiology and
Development is known.
We w ish to thank Massimo Maffei for his suggested
revisions to th e physiological chapters. This book could
not have been produced without the extensive help and
support of the outstanding team of professionals at Sinauer
Associates, now an imprint of Oxford University Press. We
are especially grateful to our project editor, Laura Green,
whose considerable expertise in plant physiology and bio-
ch emistry, and thorough dissection and critique of each
chapter, which sometimes entailed major reorgan izing,
helped us to optimize the logical flow of topics and avoid
as many factual errors as is humanly possible. Our second
line of defense against errors and stylistic problems was our
tireless copyeditor, Liz Pierson, who did wonders to shape
the newly designed chapters into a seamless, integrated, and
coherent whole. As always, the talented artist Elizabeth Mo-
rales implemented all the changes we requested to existing
figures an d created several new, attractive figures as well.
We owe the new single- column format of the book,
including the running glossary, to the creative talents of
Chris Small and his production team. Ann Chiara designed
the in side of t he book an d crafted the layout. Another
new design feature was use of ch apter openers featuring
plants pertinent to each chapter's content. Mark Siddall,
our photo researcher, is responsible for finding the many
fine and appropriate photos for u se as chapter openers, and
Beth Roberge Friedrich s deserves credit for design ing the
distinctive new cover for Fundamentals of Plant Physology.
We are also grateful to Johann a Walkowicz for laying the
foundation for the book's conception by devising and com-
piling surveys obtained from a wide selection of instructors.
Thanks to her detailed an alysis of the data, we were able
to identify those topics of plant physiology that were most
often covered in courses.
Finall:Yt we extend our deepest gratitude to our publisher,
Andy Sinauer, whose creative and innovative contribution s
to Fundamentals of Plant Physiology, as well as to previous
editions of Plant Physiology and Development, are too nu mer-
ous to list. Andy is retiring this year, and we are extremely
sorry to see him go. He leaves behind a magnificent legacy,
Sinauer Associates, that is second to none among publishers
of scientific textbooks. Although we will sorely miss Andy's
w isdom and invaluable insights, we look forward to a
productive ongoing relationship with our new publisher,
Oxford Un iversity Press.
Lincoln Taiz Eduardo Zeiger
Ian Max Moller Angus Murphy
April 2018
E= itors
Lincoln Taiz is Professor Emeritus of
Molecular, Cellular, and Developmental
Biology at the University of California at
Santa Cruz. He received his Ph.D. in Bot-
any from the University of California at
Berkeley in 1971. Dr. Taiz's main research
focus has been on the structure, function,
and evolution of vacuolar H+-ATPases. He
has also worked on gibberellins, cell wall
mechanical properties, metal tolerance,
auxin transport, and stomatal opening.
lan Max M0ller is Professor Emeritus of
Molecular Biology and Genetics at Aar-
hus University Denmark. He received his
Ph.D. in Plant Biochemistry from Imperial
College, London, UK. H e has worked at
Lund University, Sweden and, more recently,
at Ris0 National Laboratory and the Royal
Veterinary and Agricultural University in
Copenhagen, Denmark. Professor M0ller
has investigated plant respiration throughout his career. H is
current interests include turnover of reactive oxygen species and
the role of protein oxidation in plant cells.
Contributors
Sarah M. Assmann Susan Dunford
Christine Beveridge James Ehleringer
Robert E. Blankenship Jurgen Engelberth
Arnold J. Bloom Lawrence Griffing
Eduardo Blumwald N. Michele Holbrook
John Browse Andreas Madlung
Bob B. Buchanan Ron Mittler
Victor Busov Gabriele B. Monshausen
John Christie Wendy Peer
Daniel J. Cosgrove Allan G. Rasmusson
Eduardo Zeiger is Professor
Emeritus of Biology at the University
of California at Los Angeles. He
received a Ph.D. in Plant Genetics
at the University of California at
Davis in 1970. H is research inter-
ests include stomatal function, the
sensory transduction of blue-light
responses, and the study of sto-
matal acclimations associated with
increases in crop yields.
Angus Murphy has been a Professor and
Chair of the Department of Plant Science
and Landscape Architecture at the Univer-
sity of Maryland since 2012. H e earned his
Ph.D. in Biology from the University of Cal-
ifornia at Santa Cruz in 1996 and moved to
Purdue University as an assistant professor
in 2001. Dr. Murphy studies ATP-Binding
Cassette transporters, the regulation of
auxin transport, and the mechanisms by which transport pro-
teins are regulated in plastic plant growth.
Darren R. Sandquist
Graham B. Seymour
Sally Smith
Joe H. Sullivan
Heven Sze
Bruce Veit
Philip A. Wigge
Ricardo A. Wolosiuk
to accompany Fundamentals of Plant Physiology
eBook
(ISBN 978-1-60535-791-1)
Fundamentals of Plant Physiology is available as an
eBook, in several different formats, including RedShelt
VitalSource, and Chegg. All major mobile devices
are supported.
For the Instructor
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Includes the following resources to aid instructors in
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reformatted for optimal readability, with complex
figures provided in both whole an d split formats.
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 Rem i Masson/Biosphoto/Getty Images

Pant an __ Ce Arc itecture

lant physiology is the study of plant processes-how plants grow,

develop, and function as t hey int eract w ith their physica l (abiotic) and
living (biotic) environments. A lthoug h this book w ill emphasize the
physiological and biochemical functions of plants, it is import ant to recog-
nize that, w hether we are ta lking about gas exchange in the leaf, wat er con-
duction in t he xylem, photosynthesis in the chloroplast, ion transport across
membranes, signal transduction pathways involving light and hormones, or
gene expression during development, all of these physiologica l and bio-
chem ica l functions depend entirely on structures. Function derives from
structures interacting at every level of scale, from molecules to organisms.
The cell is t he fundamental organizationa l unit of plants and all other
living organisms. The term cell is derived from the Latin cella, meaning
11 11
IIstoreroom or Chamber. It was first used in biology in 1665 by the
II

English scientist Robert Hooke to describe the individual units of the honey-
comb-like structure he observed in cork under a compound microscope. The
cork llcellsll Hooke observed were actually the empty lumens of dead cells
surrounded by cell walls, but the term is an apt one, because cells are the
basic building blocks that define plant structure.
Groups of special ized cells form specific tissues, and specific tissues
arranged in particular patterns are the basis of three-dimensional o rgans.
Plant anatomy is the study of the macroscopic arrangements of cells and
tissues with in organs, and plant cell biology is the st udy of th e organelles
and other small components that make up each cell. Improvements in both
light and electron microscopy continue to reveal th e ast onish ing va riety
and dynamics of cel lular processes and their contributions to physiolog ica l
functions .
2 Chapter 1

This chapter provides an overview of the basic anatomy and cell biology of
plants, from the macroscopic structure of organ s and tissues to the m icroscopic
ultrastructure of cellular organelles. Subsequent chapters will treat these structures
in greater detail from the perspective of their physiological and developmental
functions at d ifferen t stages of the plant life cycle.

Plant Life Processes: Unifying Principles

The spectacular diversity of plant size and for m is familiar to everyone. Plants
range in height from less than 1 em to more than 100 m . Plant morphology, or
form, is also surprisingly diverse. At first glance, the tiny plant duckweed (Lemna)
seems to have little in common with a giant saguaro cactus or a redwood tree. No
single plant shows the entire spectrum of environmental adaptations that allow
plants to occupy almost every n iche on Earth, so plant physiologists often study
model organisms that are representative of major plant fu nctions and are easy
to manipulate in research stud ies. These model systems are useful because all
plants, regardless of their specific adaptations, carry out fu ndamentally similar
processes and are based on the same architectural plan.
We can summarize the major unifying principles of plants as follows:
• As Earth's primary producers, plants and algae are the ultimate solar
collectors. They harvest the energy of sunlight by converting light en-
ergy to chemical energy, which they store in bonds formed when they
synthesize carbohydrates from carbon dioxide and water.
• Except for certain reproductive cells, plants do not move from place to
place; they are sessile. As a substitute for motility, they have evolved the
ability to grow toward essential resources, such as light, water, and m in-
eral nutrients, throughout their life span.
• Plants are structurally reinforced to support their mass as they grow to -
ward sunlight against the pull of gravity.
• Plants have mechanisms for moving water and minerals from the soil
to the sites of photosynthesis and growth, as well as mechanisms for
moving the products of photosynthesis to nonphotosynthetic organs
and tissues.
• Plants lose water continuously by evaporation and h ave evolved mech a-
nisms for avoiding desiccation.
• Plants develop from embryos that derive nutrients from the mother
plant, and these add itional food stores facilitate the production of large
self-supporting structures on land.

Plant Classification and Life Cycles

Based on the principles listed above, we can define plants generally as sessile,
multicellular organisms derived from embryos, adapted to land, and able to
convert carbon dioxide into complex organic compounds th rough the process of
photosynthesis. This broad definition includes a wide spectrum of organisms,
from the mosses to the flowering plants (Box 1.1 ).
With the exception of the great coniferous forests of Can ada, Alaska, and
northern Eurasia, angiosperm s dominate the landscape today. More than 250,000
species are k nown, w ith tens of thousands of additional undescribed species pre-
d icted by computer models. Many of the predicted species are imperiled because
they occur primarily in regions of rich biodiversity where habitat destruction is
common. The major an atomical innovation of the angiosperms is the flower;
 Plant and Cel l Architecture 3

Box 1.1 Evolutionary Relationships among Plants

Plants share w ith (mostly aquatic) green algae t he primitive
trait t hat is so important for photosynthesis in both clades:
Their chloroplasts contain the pigments ch lorophyl l a
and band ~-carotene . Plants, or embryophytes, share
the evolutionarily derived traits for surviving on land that
are absent in the algae (see f igure). Plants include the
nonvascular plants, referred to as bryophytes (mosses,
hornworts, and liverworts), and the vascular plants, or
tracheophytes . The vascu lar plants, in turn, consist of the
non-seed plants (ferns and thei r relatives) and the seed
plants (gymnosperms and ang iosperms).
Because plants have many ag ricu ltural, industrial,
timber, and med ical uses, as well as an overwhelm ing
dom inance in terrestrial ecosystems, most research in plant
biology has focused on the plants that have evolved in
the last 300 m illion years, the seed plants (see f igu re). The
gymnosperms (from the Greek for "naked seed") include
the conif ers, cycads, ginkgo, and gnetophytes (which
include Ephedra, a popular med icinal plant). About 800
species of gymnosperms are known. The largest group
of gymnosperms is the conifers ("cone-bearers"), which
include such commercial ly important forest trees as pine, f ir,
spruce, and redwood. The evolution of the angiosperms
(from the Greek for "vessel seed"), remains, in the words
of Darwin, an "abominable mystery." The earl iest known
ang iosperm fossi l dates to the early Cretaceous, -125
million years ago. However, based on analyses of the DNA
sequences of extant angiosperms, the basa l (i.e., prim itive)
ang iosperms (Amborellales, Nyphaeles, and Austrobaileya les)
may have appeared much earlier, in the Middle to Late
Triassic period, -224-240 m il lion years ago. The same study
suggests that the diversification of the angiosperms into
the monocots, magnoliids, and eud icots probably occurred
du ring the Jurassic period, -154-191 mill ion years ago,
around the time when important pollinating insects, such as
bees and butterflies, were also evolving. If these estimates
are correct, fossi ls corresponding to these earlier dates may
turn up in the future.

Plants
(embryophytes)

Nonvascular plants Vascular plants

Non-plants (bryophytes) (tracheophytes)
...--~A...__~
( \
Seed plants

Flowering plants (angiosperms)

Mosses,
hornworts, Ferns, Basal Magnolia
Red algae Green algae liverworts fern allies Gymnosperms ang 1osperms Monocots family Eudicots
'"7
sst
'\. 7.
~'..Yc;

'-.?
6's~
c; /
.'

%,c;/ .
'"~Oo
~0

I
1/

%,c;/ • Flowers

st,s.0
_y~
'0
%,c;/ .
I
I
seeds

~/ • Water and photosynthate

7~
j vascular transport
Oo
~c;
• Land-dwell ing
adaptations
I
Cladogram showing the evolutiona ry relationships
•Chloroplasts contain ing
among the various members of the plants and chlorophyll a+b
their close relatives, the algae. The sequence of
evolutionary innovations given on the right side of
the figure eventua lly gave rise to the angiosperms.
(Mya, million years ago.)
4 Chapter 1
alternation of generations
The presence of two genetica lly distinct
m ulticellular stages, one haploid and
one diploid, in the plant life cycle. The
haploid gametophyte generation beg ins
with meiosis, whi le the diploid sporo-
phyte generation beg ins with the fusion
of sperm and egg .
gamete A haploid (1 N) reproductive
cell.
meiosis The reduction division
II II
whereby two successive cell divisions
produce four haploid (1 N) cells from one
diploid (2N) cel l. In plants with alterna-
t ion of generations, spores are produced
by meiosis. In animals, wh ich don't have
alternation of generations, gametes are
produced by meiosis .
spores Reproductive cells formed
in plants by meiosis in the sporophyte
generation. They give rise by mitotic divi-
sions to the gametophyte generation .
sporophyte The diploid (2N) multi-
cellula r structure t hat produces haploid
spores by meiosis.
mitosis The ordered cellular process
by wh ich replicated chromosomes are
distributed to daughter cells formed by
cytokinesis .
gametophyte The haploid (1 N) mul-
t icellula r structu re t hat produces haploid
gametes by mitosis and differentiation .
pollen Small structures (microspores)
produced by anthers of seed plants.
Conta in haploid male nuclei that will
fertilize the egg in the ovule.
fertilization The formation of a
diploid (2N) zygote from the cellular
and nuclear fusion of two haploid (1 N)
gametes, the egg and the sperm. In
angiosperms, ferti lization also involves
fusion of a second sperm nucleus w ith
t he haplo id nuclei (usually two) of the
centra l cell to form the endosperm (usu-
ally triploid).
megaspore The haploid (1N)
spore t hat develops into t he female
gametophyte.
micros pores The haploid (1 N) cell
t hat develops into t he pollen tube or
male gametophyte.
monoecious Refers to plants in which
male and fe male flowe rs are found on
t he same individua ls, such as cucumber
(Cucumis sativus) and maize (corn; Zea
mays).
hence they are referred to as flowering plants, distinguished from gymnosperms
by the presence of a carpel that encloses the seeds.
Plant life cycles alternate between diploid and haploid
generations
Plants, unlike animals, alternate between two distinct multicellular generations to
complete their life cycle, a distinctive feature termed alternation of generations.
One generation has diploid cells, cells with two copies of each chromosome and
abbreviated as having 2N chromosomes, and the other generation has haploid
cells, cells w ith only one copy of each ch romosome, abbreviated as l N. Each of
these multicellular generations may be more or less physically dependent on the
other, depending on their evolutionary grouping.
W hen diploid (2N) animals such as humans produce haploid gametes, egg
(lN) and sperm (lN), they do so directly by the process of meiosis, cell division
resulting in a reduction of the number of chromosomes from 2N to l N. This
cycle is depicted in the in ner portion of Figure 1.1 . In contrast, the products of
meiosis in diploid plants are spores, and diploid plant forms are therefore called
sporophytes. Each spore is capable of u ndergoing mitosis, cell division that does
not change the number of ch romosomes in the daughter cells, to form a n ew
h aploid mu lticellu lar individual, the gametophyte. These cycles are depicted in
the outer portion of Figure 1.1. The h aploid gametophytes produce gametes, egg
and sperm, by simple mitosis, whereas haploid gametes in animals are produced
by meiosis. This is a fu ndamental difference between plants and animals and
gives the lie to some stories about "the birds and the bees"-bees do not carry
around sperm to fertilize female flowers, they carry arou nd the male gametophyte,
the pollen, which is a multicellular structure that produces sperm cells. When
placed on receptive sporophytic tissue, the pollen grain germinates to form a
pollen tube that must grow through sporophytic tissue u ntil it reaches the female
gametophyte. The male gametophyte penetrates the female gametophyte and
releases sperm to fertilize the egg.
O nce the haploid gametes fuse and fertilization takes place to create the 2N
zygote, the life cycles of animals and plants are similar (see Figure 1.1). The 2N
zygote undergoes a series of mitotic division s to produce the embryo, wh ich
eventually grows into the mature diploid adult.
Thu s, all plant life cycles encompass two separate generations: the diploid,
spore-producin g sporophyte generation an d the haploid, gamete-producing
gametophyte generation . A line drawn between fertilization and meiosis divides
these two separate stages of the generalized plant life cycle shown in Figure 1.1.
Increasing the nu mber of mitoses between fertilization and meiosis increases the
size of the sporophyte generation and the number of spores that can be produced.
Having more spores per fertilization event could compen sate for low fertility
when water becomes scarce on land. This cou ld explain the marked tendency
for the increase in size of the sporophyte generation, relative to the gametophyte
generation, du ring the evolution of plants.
The sporophyte generation is dominant in the seed plants (gymnosperms and
angiosperms) and gives rise to the megaspores, wh ich develop into the female
gametophyte, and the microspores, which develop into the male gametophyte (see
Figure 1.1). Both megaspores and microspores produce gametophytes with relatively
few cells compared with the sporophyte. In the vast majority of angiosperms,
both male and female gametophytes occur in a single hermaphroditic ("perfect")
flower, as in tulips. In some angiosperms, the male and female gametophytes
occur on different flowers on the same plant, as in the male (staminate) and
female (pistillate) flowers of maize (corn; Zea mays). Such species are referred
to as monoecious (from the Greek for "one hou se"). Alternatively, if the male
and female flowers occur on separate individuals, as in spinach plants or willow
 Plant and Cel l A rchitecture 5

SPOROPHYTE GEN ERATION GAMETOPHYTE GENERATION

o Megaspo re (1N) <;j? Ga metophyt e (1 N) =embryo sac

~ .. Meiosis 0 Microspo re (1N) M itosis o Gametophyte (1 N) = pol len

Spore (1N)

Gamet ophyte (1N)

with o
and <;j? o rgans
(antheridium and archegoni um)

o and <;j? o and <;j? Sorus Capsule

flowers cones Meiosis

l Moss Fern

Human
Ang iosperm Gymnosperm Fern Moss
Sperm Egg Sperm Egg Sperm Egg
Sporophyt e (2N) Sporophyt e (2N) grows out (1N) (1N) (1N) (1N) (1N) (1N)
of gametophyte (1N)
No seed
Germination

Ferti Iizati on

Zygot e Zygote Zygote

Seed with embryo (2N) (2N) (2N) (2N)
Seed coat f rom sporophyt e
Embryo (2N)

.......__ _ _ _ _ _ _ _ __Mitosis - - - - - - - - - '

Embryo (2N) w ithin
archegonium (1N)

Embryo (2N) within embryo sac (1N in

gymnosperms, 3N double-fertilized
endosperm in angiosperms)

Figure 1.1 Diagram of the generalized life cycles of plants and animals. In contrast
to animals, plants exhibit alternation of generations. Rather than producing gametes
directly by meiosis as an ima ls do, plants produce vegetative spores by meiosis. These
1N (haplo id) spores divide to produce a second multicellular ind ividual called the game-
tophyte. The gametophyte then produces gametes (sperm and egg) by m itosis. Follow-
ing fertil ization, the resulting 2N (d iploid) zygote develops into the mature sporophyte
generation, and the cycle beg ins again. In ang iosperms, the process of double fertili-
zation produces a 3N (triploid) or higher ploidy level (see Chapter 17) feed ing tissue
called the endosperm.
6 Chapter 1
dioecious Refers to plants in wh ich
male and female flowers are found on
different individuals, such as spinach
(Spinacia) and hemp (Cannabis sativa) .
double ferti lization A un ique fea-
ture of all angiosperms whereby, along
with the fusion of a sperm with the egg
to create a diploid zygote, a second
male gamete fuses with the polar nuclei
in the embryo sac to generate the endo-
sperm tissue (with a triploid or higher
number of chromosomes).
stem The typically above ground pri-
mary axis of the shoot that bears leaves
and buds. May also occur underground
in the form of rhizomes, corms, and
tubers.
root The organ, usual ly underground,
that serves to anchor the plant in the
soil, and to absorb water and mineral
ions, and conduct them to the shoot.
In contrast to shoots, roots lack buds,
leaves, or nodes.
leaves The main latera l appendages
rad iating out from stems and branches.
Green leaves are usua lly the major pho-
tosynthetic organs of the plant.
Soil line
Figure 1.2 Schematic representation
of the body of a typica l eud icot.
trees, the species is termed dioecious (from the Greek for "two houses"). In
gymnosperms, ginkos and cycads are dioecious, while conifers are monoecious.
Con ifers produce female cones, called megastrobili, w h ich are u sually higher
up on the tree, an d male con es, called microstrobili, on the lower branch es, a n
arrangement th at in creases the chances of cross-pollin ation with other trees.
Sperm and egg production, as well as the dyn amics of fertilization, differs
among gametophytes of seed plants. Angiosperms carry out the u n ique process
of double ferti lization, in which two sperm cells are produced, only one of wh ich
fertilizes the egg. Th e other sperm fuses w ith two nu clei in the female gametophyte
to produ ce the 3N (th ree sets of ch romosomes) endosperm, the storage tissue for
the angiosperm seed. (Some angiosperms produce endosperm of h igher ploidy
levels; see Ch apter 17.) Th e storage tissue for th e seed in gymn osperms is 1N
gametophytic tissue because there is no double fertilization (see Figure 1.1). So
the seed of seed plants is n ot at all a spore (defin ed as a cell that produ ces th e
gametophyte generation), but it does contain gametophytic (1N) storage tissue
in gymnosperms an d gametophy te -derived 3N storage tissue in an giosperms.
In the lower plants su ch as fern s and mosses, the sporophyte generation
gives rise to spores th at grow into adult gametophytes th at then h ave regions
that differentiate into male an d female structures, the male antheridium and the
female archegon ium. In ferns the gametophyte is a small monoecious prothallu s,
which has antheridia a n d arch egonia that divide m itotically to p roduce motile
sperm and egg cells, respectively. Th e dominant leafy gametophyte generation
in mosses contains an th erid ia an d archegonia on the same (monoecious) or
different (dioecious) individuals. Th e motile sperm then enters the archegonium
and fertilizes th e egg, to form the 2N zygote, w h ich develops into an embryo
enclosed in th e gametophy tic tissue, bu t no seed is formed. The embryo directly
develops into the adult 2N sporophyte.
Overview of Plant Structure
Despite their apparent diversity, all seed plants have the same basic body plan
(Figure 1.2). The vegetative body is composed of th ree organs-the stem, th e
root, and the leaves-each with a different direction, or polarity, of growth. Th e
stem grows upward and supports the aboveground part of the plant. The root,
which anchors the plant and absorbs nutrients a n d water, grows down below
the groun d . The leaves, whose primary function is photosynth esis, grow out
laterally from the stem at the nodes. Variation s in leaf arrangement can give rise
to many different forms of shoots, th e term for the leaves a n d stem together.
For example, leaf nodes can sp iral aroun d th e stem, rotatin g by a fixed angle
between each internode (the region between two nodes). Altern ativelY" leaves
can arise oppositely or altern ating on eith er side of th e stem.
O rgan shape is defined by directional p attern s of growth . Th e p olarity of
growth of the primary plant axis (th e main stem and root) is vertical, whereas
the typical leaf grows laterally at the margins to produce the flattened leaf blade.
The growth polarities of these organ s are adapted to their fun ctions: Leaves
fun ction in light absorption, stems elongate to lift the leaves toward sun light,
and roots elongate in search of water and nutrients from the soil. The cellular
component that directly determines growth polarity in plants is th e cell wall.
Plant cells are surrounded by rigid cell walls
The outer boundary of the living cytoplasm of plant cells is the plasma mem-
brane (also called the plasmalemma), similar to the situation in animals, fun gi,
and bacteria. Th e plasma membrane surrounds th e cytoplasm, which consists
of organelles and su pporting elements suspen ded in an aqueous cytosol. (Th e
nucleus, which is surrou nded by a double membrane a n d contain s the ch romo-
 Plant and Cell A rchitect ure 7

Compound middle lamel la

(primary walls and middle lamel la)

Nucleus Cytoplasm

Pr imary wall node Posit ion on t he stem where

Secondary wall leaves are attached .

Plasma membrane shoots The organ, usually abo-

veground, that includes the stem, leaves,
buds, and reproduct ive structures. Func-
Figure 1.3 Primary an d secondary cel l walls and t heir re lat ionsh ip t o the rest of the
t ion in photosynthesis and reproduction .
cell. The two adjacent primary wal ls, along with the m iddle lamella, form a composite
struct ure called t he compoun d middle lamella. primary plant axis The longitud inal
axis of t he plant defined by the positions
of the shoot and root apica l meristems.
leaf blade The broad, expanded area
somes, is con sidered separate from the cytoplasm.) Outside the plasma mem- of the leaf; also cal led the lamina.
brane of plant cells is a rigid cell wall composed of cellu lose and other polymers
that add rigidity and strength (Figure 1.3). During an imal development cells plasma membrane (plasma-
are able to migrate from one location to another, and tissues may thus contain lemma) A bilayer of polar lipids
(phospholipids or glycosylglycerides) and
cells that origin ated in different parts of the organ ism. In plants, however, cell embedded proteins that together form a
walls limit cellular migration, and development depends solely on patterns of selectively permeable boundary around
cell division and enlargement. a cell .
Plant cells have two types of walls: primary and secondary (see Figure 1.3). cytoplasm The cellular matter
Primary cell walls are typically th in (less than 1 ~-tm) and are characteristic of enclosed by the plasma membrane
young, growing cells. Secondary cel l walls are thicker and stronger than primary exclusive of the nucleus.
walls and are deposited on the in ner surface of the prim ary wall after most cell cytosol The aqueous phase of t he
enlargement has ended. The cell walls of n eighboring cells are cemented by cytoplasm conta ining dissolved solutes
a middle lamella composed of the carbohydrate polymer pectin. Because it is but excluding supramolecular structures,
sometimes difficult to distinguish the middle lamella from the primary walt such as ribosomes and components of
the cytoskeleton .
particularly when a secondary wall is present, the two adjacent primary walls
and middle lamella are termed the compound m iddle lamella. cell wall The rigid cel l surface struc-
ture externa l to t he plasma membrane
Primary and secondary cell walls differ in their components that supports, binds, and protects the
cell. Composed of cellulose and other
Cell walls contain various typ es of polysaccharides that are n amed after the polysaccharides and proteins.
principal su gars th ey contain . For example, a glucan is a p olymer of glucose
units linked end to end. Polysaccharides may be linear u nbranched chain s of primary cell walls The thin (less than
1 ~m), unspecialized cell walls that are
sugar residues, or they may contain side branches attached to the backbone. For characteristic of young, growing cells.
branched polysaccharides, the backbone of the polysaccharide is usu ally indicat-
ed by the last part of the n ame. For example, xyloglucan has a glucan backbone secondary cell wall Cell wa ll synthe-
sized by nongrowing cells. Often multi-
(a linear chain of glucose residues) with xylose sugars attached as side chains. layered and contain ing lign in, it differs
The specific lin kages between sugar rings, including the specific carbons that in composit ion and struct ure from the
are linked together and the configuration of the lin kage, are important for the primary wa ll.
properties of the polysacch aride. For instance, amylose (a component of starch in middle lamella A thin layer of pec-
the plastid) is an a(1,4)-linked glucan (C-1 and C-4 carbon s of adjacent glucose t in-rich mat eria l at the junction where
rings are linked th rough an 0 -glycosidic bond in an a con figuration), w hereas the primary walls of neighboring cells
cellu lose is a glucan made of ~(1,4) -linkages (Figure 1.4). These differences in come into contact.
linkages m ake huge differences in the physical properties, enzymatic digestibility glucan A polysaccharide made from
and functional roles of these two polymers of glucose. glucose un its.
8 Chapter 1

a-1 ,4 Glycosidic linkage

A
6 CH 20H CH20H CH20 H ,-------- ----.., CH20 H
H 5 H H OH Formation H A-----0 H H 0 OH
0 0
~ of a linkage ~
4 1 + 4 1 )II 4 1 1

OH 2
a
mD mo H
\ 0
H
H
3
OH H OH
E H OH H OH
o.-o-G Iucose 0-o-G Iucose Further
polymerizat ion

CH20H CH20H
H H H H 0 0

"o 0 0 0 0/

H H
Amylose

~-1 ,4 Glycosidic lin kage

A
6 CH 20H CH 0 H ,----- ----.., H OH
2
·J ---0 mD H OH Formation H J---0
0
H
~ ~ of~ linkage 1 ~
4

OH ~3 -~ H
1 + 4

mo
1
\. OH
4
1

OH
H H OH E H OH
~-o-Giucose ~-o-G Iucose Further
polymerizat ion

H H
H
0 0
"o 0

H
Cellulose

Figure 1.4 Structures of amylose and cellu lose. The hydroxyl groups of amylose and
cellu lose are high lighted in blue to emphasize the structural differences between the
polymers.

cellulose A linear cha in of (1 A )- Cell wall polysaccharides are classified into three groups: cellulose, hem i-
linked ~ -o-g l ucose. The repeating unit is celluloses, and p ectin. Cellulose is the major fibrillar component of the cell
cellobiose.
wall and is composed of an array of ~(1,4) -linked glucans aggregated to form a
microfibril The major fibrillar com- microfibril with well-ordered and less-ordered region s. The simplest cellulose
ponent of the cell wal l composed of microfibrils are narrow structures, approxim ately 3 n m wide (1 n m =10-9 m),
layers of cel lulose molecules packed
which reinforce the cell wall like the steel rods in reinforced con crete. Each
t ightly together by extensive hydrogen
bonding. microfibril is composed of an estim ated 18 parallel ch ain s of (1,4) -linked
~ - D -glucose tightly packed together to form a crystallin e core with extensive
hemicelluloses Heterogeneous group
of polysaccharides that bind to the sur-
hydrogen bonding within and between glucan chain s (Figure 1.5). Microfibrils
face of cellulose, linking cel lulose m icro- are insoluble in water and have a h igh tensile stren gth, about half the tensile
f ibri ls together into a network. strength of steel.
Hemicelluloses are a heterogeneous group of polysacch arides w ith ~ (1,4) ­
pectins A heterogeneous group of
complex cell wall polysaccharides that linked backbones. Hemicellu loses typically requ ire a strong reagent, such as 1- 4
form a gel in which the cel lulose-hem i- M NaOH, to be extracted from the cell wall. The third group of major cell wall
cellulose net work is embedded. components is pectin, a diverse group of hydrophilic, gel-forming polysacch a-
 Pl ant and Cell A rchitect ure 9

(A) (C) Cross sectio n of cell u lose m icrof ibri l

Crystal l ine core

is h ighly o r gan ized

Su rf ace g l u ca n s
a re less ord er ed

(D) Hy d rogen b o n d ing between a n d w ith in

adjacen t g luca n chai n s
(B) ••
0 •• 0 •
: :
H H ••:
0 0 0

OH ...• O 9H ·····o OH
;
•
H 0 •• H 0 •
••
H
•
·.•. H
0 0 0

500 n m HO ..HO·.
-.• 0 ...
•• 0 ·····
•• 0 H ••. 0
Single cell ulose m icrof ibril •.
H H
0 0

Figure 1.5 Structure of a cellulose microfibril. (A) Atomic force microscopy image
of the primary cell wa ll from onion epidermis. Note its fibril lar text ure, which arises
from layers of cellulose m icrofibri ls. (B) A single cellulose m icrofibril composed of
(1 A)- ~-o-glucan chains tightly bonded t o each other to form a crystall ine microfibril.
(C) Cross section of a cel lulose microfibri l, il lustrat ing one model of cellulose structure,
w ith a crystal line core of highly ordered (1 A)- ~-o-glucans surrounded by a less organ-
ized layer. (D) The crystall ine regions of cellulose have precise al ignment of glucans,
w ith hydrogen bonding (indicated by dotted red lines) within, but not between, layers
of (1A)-~-o-g l ucans . (After Matthews et al. 2006; micrograph from Zhang et al. 20 13.)

rides rich in acidic sugar residues, such as galacturonic acid. Many pectins are
readily solubilized from the wall with hot water or with Ca2+ chelators. Together,
hemicelluloses and pectins make up the matrix polysaccharides.
Typical primary cell walls of eudicots are rich in pectins, with smaller amounts
of cellulose and hemicelluloses, w hile secondary cell walls are high in cellulose
and a different form of hemicellulose. Instead of pectin, secondary walls have
varying amounts of lignin, a highly cross-lin ked polymer of aromatic alcohols
that confers rigidity to secondary walls. Whereas t he high pectin content of
primary cell walls en ables them to expan d during cell enlargement, the cellu-
lose-hemicellulose-lignin complex of secondary cell walls forms a rigid matrix
that is well suited for strength and compression resistance.
The evolution of lignified secondary cell walls provided plants with the struc-
tural rein forcement necessary to grow vertically above the soil and to colonize
the land . Bryophytes, which lack lignified cell walls, are u nable to grow more matrix polysaccharides Po lysaccha-
rides comprising the matrix of plant cel l
than a few centimeters above the grou nd. walls. In primary cel l walls they consist of
pectins, hem icelluloses, and proteins.
The cellulose microfibrils and matrix polymers are synthesized
via different mechanisms lignin High ly branched phenolic
polymer made up of phenylpropanoid
Cellulose microfibrils are synthesized by large, ordered protein complexes, alcohols that is deposited in secondary
called cellulose synthase (CESA) complexes, which are embedded in the plas- cell walls.
10 Chapt er 1
Figure 1.6 Cellu lose microfibrils are synthesized at the cel l
su rface by p lasma membrane-bound comp lexes contai ni ng
cellulose synthase (CESA) p roteins. Computationa l model
of a CESA complex extruding glucan cha ins that coalesce to
form a m icrofibril.
cellulose synthase Enzyme that
cata lyzes the synthesis of individual
(1 ,4)-li nked ~-o-g l uca n s that make up
cellulose m icrofibri ls.
plasmodesmata (singular p las-
m odesma) Microscopic membra ne-li ned
cha nnel connecting adj acent cells
t hrough the cell wal l and fi lled w ith
cytoplasm and a central rod derived
f rom t he ER cal led the desmotubule.
symplast The contin uous system
of cell protoplasts interconnected by
plasmodesmata.
symplastic transport The inter-
cellula r transport of water and solutes
t hrough plasmodesmata.
apoplastic transport Movement
of molecules through the cell wall
contin uum that is called the apoplast.
Molecu les may move through the li nked
cell walls of adjacent cel ls, and in that
way move throughout the pla nt without
crossing a plasma membrane.
size exclusion limit (SEL) The
restriction on the size of molecules that
can be transported via the symplast. It is
imposed by the w idth of the cytoplasmic
sleeve that surrounds the desmotubule
in the center of the p lasmadesma.
M icrof ilbril
Outsid e of cel l
Plasm a
Cytop lasm
CESA co m p lex
rna membrane (Figure 1.6). These rosette-like structures are made up of six
subunits, each of which is believed to contain three to six units of cellu lose
synthase, the enzyme that synthesizes the individu al glucans that make up
the microfibril. The catalytic domain of cellu lose synthase, which is located
on the cytoplasmic side of the plasma membrane, transfers a glucose residue
from a sugar nucleotide donor, uridine diphosphate glucose (UDP-glucose),
to the growing glucan chain.
In contrast to the cellulose microfibrils, the matrix polysaccharides are syn-
thesized by membrane-bou nd enzymes that polymerize sugar molecules within
the Golgi apparatus and are delivered to the cell wall via the exocytosis of tiny
vesicles (Figure 1.7). Additional sugar residues may be added as branches to the
polysaccharide backbone by other sets of enzymes. Unlike cellulose, which forms
a crystalline microfibril, the matrix polysaccharides are much less ordered and
are often described as amorphous. This amorphous character is a consequence
of the structure of these polysaccharides-their branching and their nonlinear
conformation. The predominant hemicellulose in primary cell walls of most land
plants is xyloglucan, whereas the major hemicellulose in the primary cell wall
of grasses (Poaceae) is arabinoxylan.
Plasmodesmata allow the free movement of molecules
between cells
The cytoplasm of neighboring cells is usually connected by means of plas-
modesmata (singular plasmodesma), tubular stru ctures 40 to 50 nm in diam-
eter and formed by the connected membranes of adjacent cells (Figure 1.8).
Plasmodesmata facilitate intercellular movement of proteins, nucleic acids,
and other macromolecular signals that coordinate developmental processes
between plant cells. Plant cells interconnected in this way form a cytoplas-
mic continuum referred to as the symplast, and transport of small molecu les
through plasmodesmata is called symplastic transport (see Chapters 3 and
6). Transport through the permeable cell wall space outside the cells is called
apoplastic transport. Both forms of transport are important in the vascular
system of plants (see Chapter 6).
The symplast can transport water, solutes, and macromolecules between
cells without crossing the plasma membrane. However, there is a restriction on
the size of molecules that can be transported via the symplast; this restriction
is called the size exclusion limit, and it varies with cell type, environment, and
developmental stage. The transport can be followed by studying the movement
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 DANCE ON STILTS AT THE GIRLS’ UNYAGO, NIUCHI

Newala, too, suffers from the distance of its water-supply—at least

the Newala of to-day does; there was once another Newala in a lovely
valley at the foot of the plateau. I visited it and found scarcely a trace
of houses, only a Christian cemetery, with the graves of several
missionaries and their converts, remaining as a monument of its
former glories. But the surroundings are wonderfully beautiful. A
thick grove of splendid mango-trees closes in the weather-worn
crosses and headstones; behind them, combining the useful and the
agreeable, is a whole plantation of lemon-trees covered with ripe
fruit; not the small African kind, but a much larger and also juicier
imported variety, which drops into the hands of the passing traveller,
without calling for any exertion on his part. Old Newala is now under
the jurisdiction of the native pastor, Daudi, at Chingulungulu, who,
as I am on very friendly terms with him, allows me, as a matter of
course, the use of this lemon-grove during my stay at Newala.
 FEET MUTILATED BY THE RAVAGES OF THE “JIGGER”
(Sarcopsylla penetrans)

The water-supply of New Newala is in the bottom of the valley,

some 1,600 feet lower down. The way is not only long and fatiguing,
but the water, when we get it, is thoroughly bad. We are suffering not
only from this, but from the fact that the arrangements at Newala are
nothing short of luxurious. We have a separate kitchen—a hut built
against the boma palisade on the right of the baraza, the interior of
which is not visible from our usual position. Our two cooks were not
long in finding this out, and they consequently do—or rather neglect
to do—what they please. In any case they do not seem to be very
particular about the boiling of our drinking-water—at least I can
attribute to no other cause certain attacks of a dysenteric nature,
from which both Knudsen and I have suffered for some time. If a
man like Omari has to be left unwatched for a moment, he is capable
of anything. Besides this complaint, we are inconvenienced by the
state of our nails, which have become as hard as glass, and crack on
the slightest provocation, and I have the additional infliction of
pimples all over me. As if all this were not enough, we have also, for
the last week been waging war against the jigger, who has found his
Eldorado in the hot sand of the Makonde plateau. Our men are seen
all day long—whenever their chronic colds and the dysentery likewise
raging among them permit—occupied in removing this scourge of
Africa from their feet and trying to prevent the disastrous
consequences of its presence. It is quite common to see natives of
this place with one or two toes missing; many have lost all their toes,
or even the whole front part of the foot, so that a well-formed leg
ends in a shapeless stump. These ravages are caused by the female of
Sarcopsylla penetrans, which bores its way under the skin and there
develops an egg-sac the size of a pea. In all books on the subject, it is
stated that one’s attention is called to the presence of this parasite by
an intolerable itching. This agrees very well with my experience, so
far as the softer parts of the sole, the spaces between and under the
toes, and the side of the foot are concerned, but if the creature
penetrates through the harder parts of the heel or ball of the foot, it
may escape even the most careful search till it has reached maturity.
Then there is no time to be lost, if the horrible ulceration, of which
we see cases by the dozen every day, is to be prevented. It is much
easier, by the way, to discover the insect on the white skin of a
European than on that of a native, on which the dark speck scarcely
shows. The four or five jiggers which, in spite of the fact that I
constantly wore high laced boots, chose my feet to settle in, were
taken out for me by the all-accomplished Knudsen, after which I
thought it advisable to wash out the cavities with corrosive
sublimate. The natives have a different sort of disinfectant—they fill
the hole with scraped roots. In a tiny Makua village on the slope of
the plateau south of Newala, we saw an old woman who had filled all
the spaces under her toe-nails with powdered roots by way of
prophylactic treatment. What will be the result, if any, who can say?
The rest of the many trifling ills which trouble our existence are
really more comic than serious. In the absence of anything else to
smoke, Knudsen and I at last opened a box of cigars procured from
the Indian store-keeper at Lindi, and tried them, with the most
distressing results. Whether they contain opium or some other
narcotic, neither of us can say, but after the tenth puff we were both
“off,” three-quarters stupefied and unspeakably wretched. Slowly we
recovered—and what happened next? Half-an-hour later we were
once more smoking these poisonous concoctions—so insatiable is the
craving for tobacco in the tropics.
Even my present attacks of fever scarcely deserve to be taken
seriously. I have had no less than three here at Newala, all of which
have run their course in an incredibly short time. In the early
afternoon, I am busy with my old natives, asking questions and
making notes. The strong midday coffee has stimulated my spirits to
an extraordinary degree, the brain is active and vigorous, and work
progresses rapidly, while a pleasant warmth pervades the whole
body. Suddenly this gives place to a violent chill, forcing me to put on
my overcoat, though it is only half-past three and the afternoon sun
is at its hottest. Now the brain no longer works with such acuteness
and logical precision; more especially does it fail me in trying to
establish the syntax of the difficult Makua language on which I have
ventured, as if I had not enough to do without it. Under the
circumstances it seems advisable to take my temperature, and I do
so, to save trouble, without leaving my seat, and while going on with
my work. On examination, I find it to be 101·48°. My tutors are
abruptly dismissed and my bed set up in the baraza; a few minutes
later I am in it and treating myself internally with hot water and
lemon-juice.
Three hours later, the thermometer marks nearly 104°, and I make
them carry me back into the tent, bed and all, as I am now perspiring
heavily, and exposure to the cold wind just beginning to blow might
mean a fatal chill. I lie still for a little while, and then find, to my
great relief, that the temperature is not rising, but rather falling. This
is about 7.30 p.m. At 8 p.m. I find, to my unbounded astonishment,
that it has fallen below 98·6°, and I feel perfectly well. I read for an
hour or two, and could very well enjoy a smoke, if I had the
wherewithal—Indian cigars being out of the question.
Having no medical training, I am at a loss to account for this state
of things. It is impossible that these transitory attacks of high fever
should be malarial; it seems more probable that they are due to a
kind of sunstroke. On consulting my note-book, I become more and
more inclined to think this is the case, for these attacks regularly
follow extreme fatigue and long exposure to strong sunshine. They at
least have the advantage of being only short interruptions to my
work, as on the following morning I am always quite fresh and fit.
My treasure of a cook is suffering from an enormous hydrocele which
makes it difficult for him to get up, and Moritz is obliged to keep in
the dark on account of his inflamed eyes. Knudsen’s cook, a raw boy
from somewhere in the bush, knows still less of cooking than Omari;
consequently Nils Knudsen himself has been promoted to the vacant
post. Finding that we had come to the end of our supplies, he began
by sending to Chingulungulu for the four sucking-pigs which we had
bought from Matola and temporarily left in his charge; and when
they came up, neatly packed in a large crate, he callously slaughtered
the biggest of them. The first joint we were thoughtless enough to
entrust for roasting to Knudsen’s mshenzi cook, and it was
consequently uneatable; but we made the rest of the animal into a
jelly which we ate with great relish after weeks of underfeeding,
consuming incredible helpings of it at both midday and evening
meals. The only drawback is a certain want of variety in the tinned
vegetables. Dr. Jäger, to whom the Geographical Commission
entrusted the provisioning of the expeditions—mine as well as his
own—because he had more time on his hands than the rest of us,
seems to have laid in a huge stock of Teltow turnips,[46] an article of
food which is all very well for occasional use, but which quickly palls
when set before one every day; and we seem to have no other tins
left. There is no help for it—we must put up with the turnips; but I
am certain that, once I am home again, I shall not touch them for ten
years to come.
Amid all these minor evils, which, after all, go to make up the
genuine flavour of Africa, there is at least one cheering touch:
Knudsen has, with the dexterity of a skilled mechanic, repaired my 9
× 12 cm. camera, at least so far that I can use it with a little care.
How, in the absence of finger-nails, he was able to accomplish such a
ticklish piece of work, having no tool but a clumsy screw-driver for
taking to pieces and putting together again the complicated
mechanism of the instantaneous shutter, is still a mystery to me; but
he did it successfully. The loss of his finger-nails shows him in a light
contrasting curiously enough with the intelligence evinced by the
above operation; though, after all, it is scarcely surprising after his
ten years’ residence in the bush. One day, at Lindi, he had occasion
to wash a dog, which must have been in need of very thorough
cleansing, for the bottle handed to our friend for the purpose had an
extremely strong smell. Having performed his task in the most
conscientious manner, he perceived with some surprise that the dog
did not appear much the better for it, and was further surprised by
finding his own nails ulcerating away in the course of the next few
days. “How was I to know that carbolic acid has to be diluted?” he
mutters indignantly, from time to time, with a troubled gaze at his
mutilated finger-tips.
 Since we came to Newala we have been making excursions in all
directions through the surrounding country, in accordance with old
habit, and also because the akida Sefu did not get together the tribal
elders from whom I wanted information so speedily as he had
promised. There is, however, no harm done, as, even if seen only
from the outside, the country and people are interesting enough.
The Makonde plateau is like a large rectangular table rounded off
at the corners. Measured from the Indian Ocean to Newala, it is
about seventy-five miles long, and between the Rovuma and the
Lukuledi it averages fifty miles in breadth, so that its superficial area
is about two-thirds of that of the kingdom of Saxony. The surface,
however, is not level, but uniformly inclined from its south-western
edge to the ocean. From the upper edge, on which Newala lies, the
eye ranges for many miles east and north-east, without encountering
any obstacle, over the Makonde bush. It is a green sea, from which
here and there thick clouds of smoke rise, to show that it, too, is
inhabited by men who carry on their tillage like so many other
primitive peoples, by cutting down and burning the bush, and
manuring with the ashes. Even in the radiant light of a tropical day
such a fire is a grand sight.
Much less effective is the impression produced just now by the
great western plain as seen from the edge of the plateau. As often as
time permits, I stroll along this edge, sometimes in one direction,
sometimes in another, in the hope of finding the air clear enough to
let me enjoy the view; but I have always been disappointed.
Wherever one looks, clouds of smoke rise from the burning bush,
and the air is full of smoke and vapour. It is a pity, for under more
favourable circumstances the panorama of the whole country up to
the distant Majeje hills must be truly magnificent. It is of little use
taking photographs now, and an outline sketch gives a very poor idea
of the scenery. In one of these excursions I went out of my way to
make a personal attempt on the Makonde bush. The present edge of
the plateau is the result of a far-reaching process of destruction
through erosion and denudation. The Makonde strata are
everywhere cut into by ravines, which, though short, are hundreds of
yards in depth. In consequence of the loose stratification of these
beds, not only are the walls of these ravines nearly vertical, but their
upper end is closed by an equally steep escarpment, so that the
western edge of the Makonde plateau is hemmed in by a series of
deep, basin-like valleys. In order to get from one side of such a ravine
to the other, I cut my way through the bush with a dozen of my men.
It was a very open part, with more grass than scrub, but even so the
short stretch of less than two hundred yards was very hard work; at
the end of it the men’s calicoes were in rags and they themselves
bleeding from hundreds of scratches, while even our strong khaki
suits had not escaped scatheless.

NATIVE PATH THROUGH THE MAKONDE BUSH, NEAR

MAHUTA

I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
 The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.

MAKONDE LOCK AND KEY AT JUMBE CHAURO

 This is the general way of closing a house. The Makonde at Jumbe
Chauro, however, have a much more complicated, solid and original
one. Here, too, the door is as already described, except that there is
only one post on the inside, standing by itself about six inches from
one side of the doorway. Opposite this post is a hole in the wall just
large enough to admit a man’s arm. The door is closed inside by a
large wooden bolt passing through a hole in this post and pressing
with its free end against the door. The other end has three holes into
which fit three pegs running in vertical grooves inside the post. The
door is opened with a wooden key about a foot long, somewhat
curved and sloped off at the butt; the other end has three pegs
corresponding to the holes, in the bolt, so that, when it is thrust
through the hole in the wall and inserted into the rectangular
opening in the post, the pegs can be lifted and the bolt drawn out.[50]

MODE OF INSERTING THE KEY

With no small pride first one householder and then a second

showed me on the spot the action of this greatest invention of the
Makonde Highlands. To both with an admiring exclamation of
“Vizuri sana!” (“Very fine!”). I expressed the wish to take back these
marvels with me to Ulaya, to show the Wazungu what clever fellows
the Makonde are. Scarcely five minutes after my return to camp at
Newala, the two men came up sweating under the weight of two
heavy logs which they laid down at my feet, handing over at the same
time the keys of the fallen fortress. Arguing, logically enough, that if
the key was wanted, the lock would be wanted with it, they had taken
their axes and chopped down the posts—as it never occurred to them
to dig them out of the ground and so bring them intact. Thus I have
two badly damaged specimens, and the owners, instead of praise,
come in for a blowing-up.
The Makua huts in the environs of Newala are especially
miserable; their more than slovenly construction reminds one of the
temporary erections of the Makua at Hatia’s, though the people here
have not been concerned in a war. It must therefore be due to
congenital idleness, or else to the absence of a powerful chief. Even
the baraza at Mlipa’s, a short hour’s walk south-east of Newala,
shares in this general neglect. While public buildings in this country
are usually looked after more or less carefully, this is in evident
danger of being blown over by the first strong easterly gale. The only
attractive object in this whole district is the grave of the late chief
Mlipa. I visited it in the morning, while the sun was still trying with
partial success to break through the rolling mists, and the circular
grove of tall euphorbias, which, with a broken pot, is all that marks
the old king’s resting-place, impressed one with a touch of pathos.
Even my very materially-minded carriers seemed to feel something
of the sort, for instead of their usual ribald songs, they chanted
solemnly, as we marched on through the dense green of the Makonde
bush:—
 “We shall arrive with the great master; we stand in a row and have
no fear about getting our food and our money from the Serkali (the
Government). We are not afraid; we are going along with the great
master, the lion; we are going down to the coast and back.”
With regard to the characteristic features of the various tribes here
on the western edge of the plateau, I can arrive at no other
conclusion than the one already come to in the plain, viz., that it is
impossible for anyone but a trained anthropologist to assign any
given individual at once to his proper tribe. In fact, I think that even
an anthropological specialist, after the most careful examination,
might find it a difficult task to decide. The whole congeries of peoples
collected in the region bounded on the west by the great Central
African rift, Tanganyika and Nyasa, and on the east by the Indian
Ocean, are closely related to each other—some of their languages are
only distinguished from one another as dialects of the same speech,
and no doubt all the tribes present the same shape of skull and
structure of skeleton. Thus, surely, there can be no very striking
differences in outward appearance.
 Even did such exist, I should have no time
to concern myself with them, for day after day,
I have to see or hear, as the case may be—in
any case to grasp and record—an
extraordinary number of ethnographic
phenomena. I am almost disposed to think it
fortunate that some departments of inquiry, at
least, are barred by external circumstances.
Chief among these is the subject of iron-
working. We are apt to think of Africa as a
country where iron ore is everywhere, so to
speak, to be picked up by the roadside, and
where it would be quite surprising if the
inhabitants had not learnt to smelt the
material ready to their hand. In fact, the
knowledge of this art ranges all over the
continent, from the Kabyles in the north to the
Kafirs in the south. Here between the Rovuma
and the Lukuledi the conditions are not so
favourable. According to the statements of the
Makonde, neither ironstone nor any other
form of iron ore is known to them. They have
not therefore advanced to the art of smelting
the metal, but have hitherto bought all their
THE ANCESTRESS OF
THE MAKONDE
iron implements from neighbouring tribes.
Even in the plain the inhabitants are not much
better off. Only one man now living is said to
understand the art of smelting iron. This old fundi lives close to
Huwe, that isolated, steep-sided block of granite which rises out of
the green solitude between Masasi and Chingulungulu, and whose
jagged and splintered top meets the traveller’s eye everywhere. While
still at Masasi I wished to see this man at work, but was told that,
frightened by the rising, he had retired across the Rovuma, though
he would soon return. All subsequent inquiries as to whether the
fundi had come back met with the genuine African answer, “Bado”
(“Not yet”).
 BRAZIER

Some consolation was afforded me by a brassfounder, whom I

came across in the bush near Akundonde’s. This man is the favourite
of women, and therefore no doubt of the gods; he welds the glittering
brass rods purchased at the coast into those massive, heavy rings
which, on the wrists and ankles of the local fair ones, continually give
me fresh food for admiration. Like every decent master-craftsman he
had all his tools with him, consisting of a pair of bellows, three
crucibles and a hammer—nothing more, apparently. He was quite
willing to show his skill, and in a twinkling had fixed his bellows on
the ground. They are simply two goat-skins, taken off whole, the four
legs being closed by knots, while the upper opening, intended to
admit the air, is kept stretched by two pieces of wood. At the lower
end of the skin a smaller opening is left into which a wooden tube is
stuck. The fundi has quickly borrowed a heap of wood-embers from
the nearest hut; he then fixes the free ends of the two tubes into an
earthen pipe, and clamps them to the ground by means of a bent
piece of wood. Now he fills one of his small clay crucibles, the dross
on which shows that they have been long in use, with the yellow
material, places it in the midst of the embers, which, at present are
only faintly glimmering, and begins his work. In quick alternation
the smith’s two hands move up and down with the open ends of the
bellows; as he raises his hand he holds the slit wide open, so as to let
the air enter the skin bag unhindered. In pressing it down he closes
the bag, and the air puffs through the bamboo tube and clay pipe into
the fire, which quickly burns up. The smith, however, does not keep
on with this work, but beckons to another man, who relieves him at
the bellows, while he takes some more tools out of a large skin pouch
carried on his back. I look on in wonder as, with a smooth round
stick about the thickness of a finger, he bores a few vertical holes into
the clean sand of the soil. This should not be difficult, yet the man
seems to be taking great pains over it. Then he fastens down to the
ground, with a couple of wooden clamps, a neat little trough made by
splitting a joint of bamboo in half, so that the ends are closed by the
two knots. At last the yellow metal has attained the right consistency,
and the fundi lifts the crucible from the fire by means of two sticks
split at the end to serve as tongs. A short swift turn to the left—a
tilting of the crucible—and the molten brass, hissing and giving forth
clouds of smoke, flows first into the bamboo mould and then into the
holes in the ground.
The technique of this backwoods craftsman may not be very far
advanced, but it cannot be denied that he knows how to obtain an
adequate result by the simplest means. The ladies of highest rank in
this country—that is to say, those who can afford it, wear two kinds
of these massive brass rings, one cylindrical, the other semicircular
in section. The latter are cast in the most ingenious way in the
bamboo mould, the former in the circular hole in the sand. It is quite
a simple matter for the fundi to fit these bars to the limbs of his fair
customers; with a few light strokes of his hammer he bends the
pliable brass round arm or ankle without further inconvenience to
the wearer.
SHAPING THE POT

SMOOTHING WITH MAIZE-COB

CUTTING THE EDGE

 FINISHING THE BOTTOM

LAST SMOOTHING BEFORE

BURNING

FIRING THE BRUSH-PILE

 LIGHTING THE FARTHER SIDE OF
THE PILE

TURNING THE RED-HOT VESSEL

NYASA WOMAN MAKING POTS AT MASASI

Pottery is an art which must always and everywhere excite the
interest of the student, just because it is so intimately connected with
the development of human culture, and because its relics are one of
the principal factors in the reconstruction of our own condition in
prehistoric times. I shall always remember with pleasure the two or
three afternoons at Masasi when Salim Matola’s mother, a slightly-
built, graceful, pleasant-looking woman, explained to me with
touching patience, by means of concrete illustrations, the ceramic art
of her people. The only implements for this primitive process were a
lump of clay in her left hand, and in the right a calabash containing
the following valuables: the fragment of a maize-cob stripped of all
its grains, a smooth, oval pebble, about the size of a pigeon’s egg, a
few chips of gourd-shell, a bamboo splinter about the length of one’s
hand, a small shell, and a bunch of some herb resembling spinach.
 Nothing more. The woman scraped with the
shell a round, shallow hole in the soft, fine
sand of the soil, and, when an active young
girl had filled the calabash with water for her,
she began to knead the clay. As if by magic it
gradually assumed the shape of a rough but
already well-shaped vessel, which only wanted
a little touching up with the instruments
before mentioned. I looked out with the
MAKUA WOMAN closest attention for any indication of the use
MAKING A POT. of the potter’s wheel, in however rudimentary
SHOWS THE a form, but no—hapana (there is none). The
BEGINNINGS OF THE embryo pot stood firmly in its little
POTTER’S WHEEL
depression, and the woman walked round it in
a stooping posture, whether she was removing
small stones or similar foreign bodies with the maize-cob, smoothing
the inner or outer surface with the splinter of bamboo, or later, after
letting it dry for a day, pricking in the ornamentation with a pointed
bit of gourd-shell, or working out the bottom, or cutting the edge
with a sharp bamboo knife, or giving the last touches to the finished
vessel. This occupation of the women is infinitely toilsome, but it is
without doubt an accurate reproduction of the process in use among
our ancestors of the Neolithic and Bronze ages.
There is no doubt that the invention of pottery, an item in human
progress whose importance cannot be over-estimated, is due to
women. Rough, coarse and unfeeling, the men of the horde range
over the countryside. When the united cunning of the hunters has
succeeded in killing the game; not one of them thinks of carrying
home the spoil. A bright fire, kindled by a vigorous wielding of the
drill, is crackling beside them; the animal has been cleaned and cut
up secundum artem, and, after a slight singeing, will soon disappear
under their sharp teeth; no one all this time giving a single thought
to wife or child.
To what shifts, on the other hand, the primitive wife, and still more
the primitive mother, was put! Not even prehistoric stomachs could
endure an unvarying diet of raw food. Something or other suggested
the beneficial effect of hot water on the majority of approved but
indigestible dishes. Perhaps a neighbour had tried holding the hard
roots or tubers over the fire in a calabash filled with water—or maybe
an ostrich-egg-shell, or a hastily improvised vessel of bark. They
became much softer and more palatable than they had previously
been; but, unfortunately, the vessel could not stand the fire and got
charred on the outside. That can be remedied, thought our
ancestress, and plastered a layer of wet clay round a similar vessel.
This is an improvement; the cooking utensil remains uninjured, but
the heat of the fire has shrunk it, so that it is loose in its shell. The
next step is to detach it, so, with a firm grip and a jerk, shell and
kernel are separated, and pottery is invented. Perhaps, however, the
discovery which led to an intelligent use of the burnt-clay shell, was
made in a slightly different way. Ostrich-eggs and calabashes are not
to be found in every part of the world, but everywhere mankind has
arrived at the art of making baskets out of pliant materials, such as
bark, bast, strips of palm-leaf, supple twigs, etc. Our inventor has no
water-tight vessel provided by nature. “Never mind, let us line the
basket with clay.” This answers the purpose, but alas! the basket gets
burnt over the blazing fire, the woman watches the process of
cooking with increasing uneasiness, fearing a leak, but no leak
appears. The food, done to a turn, is eaten with peculiar relish; and
the cooking-vessel is examined, half in curiosity, half in satisfaction
at the result. The plastic clay is now hard as stone, and at the same
time looks exceedingly well, for the neat plaiting of the burnt basket
is traced all over it in a pretty pattern. Thus, simultaneously with
pottery, its ornamentation was invented.
Primitive woman has another claim to respect. It was the man,
roving abroad, who invented the art of producing fire at will, but the
woman, unable to imitate him in this, has been a Vestal from the
earliest times. Nothing gives so much trouble as the keeping alight of
the smouldering brand, and, above all, when all the men are absent
from the camp. Heavy rain-clouds gather, already the first large
drops are falling, the first gusts of the storm rage over the plain. The
little flame, a greater anxiety to the woman than her own children,
flickers unsteadily in the blast. What is to be done? A sudden thought
occurs to her, and in an instant she has constructed a primitive hut
out of strips of bark, to protect the flame against rain and wind.
This, or something very like it, was the way in which the principle
of the house was discovered; and even the most hardened misogynist
cannot fairly refuse a woman the credit of it. The protection of the
hearth-fire from the weather is the germ from which the human
dwelling was evolved. Men had little, if any share, in this forward
step, and that only at a late stage. Even at the present day, the
plastering of the housewall with clay and the manufacture of pottery
are exclusively the women’s business. These are two very significant
survivals. Our European kitchen-garden, too, is originally a woman’s
invention, and the hoe, the primitive instrument of agriculture, is,
characteristically enough, still used in this department. But the
noblest achievement which we owe to the other sex is unquestionably
the art of cookery. Roasting alone—the oldest process—is one for
which men took the hint (a very obvious one) from nature. It must
have been suggested by the scorched carcase of some animal
overtaken by the destructive forest-fires. But boiling—the process of
improving organic substances by the help of water heated to boiling-
point—is a much later discovery. It is so recent that it has not even
yet penetrated to all parts of the world. The Polynesians understand
how to steam food, that is, to cook it, neatly wrapped in leaves, in a
hole in the earth between hot stones, the air being excluded, and
(sometimes) a few drops of water sprinkled on the stones; but they
do not understand boiling.
To come back from this digression, we find that the slender Nyasa
woman has, after once more carefully examining the finished pot,
put it aside in the shade to dry. On the following day she sends me
word by her son, Salim Matola, who is always on hand, that she is
going to do the burning, and, on coming out of my house, I find her
already hard at work. She has spread on the ground a layer of very
dry sticks, about as thick as one’s thumb, has laid the pot (now of a
yellowish-grey colour) on them, and is piling brushwood round it.
My faithful Pesa mbili, the mnyampara, who has been standing by,
most obligingly, with a lighted stick, now hands it to her. Both of
them, blowing steadily, light the pile on the lee side, and, when the
flame begins to catch, on the weather side also. Soon the whole is in a
blaze, but the dry fuel is quickly consumed and the fire dies down, so
that we see the red-hot vessel rising from the ashes. The woman
turns it continually with a long stick, sometimes one way and
sometimes another, so that it may be evenly heated all over. In
twenty minutes she rolls it out of the ash-heap, takes up the bundle
of spinach, which has been lying for two days in a jar of water, and
sprinkles the red-hot clay with it. The places where the drops fall are
marked by black spots on the uniform reddish-brown surface. With a
sigh of relief, and with visible satisfaction, the woman rises to an
erect position; she is standing just in a line between me and the fire,
from which a cloud of smoke is just rising: I press the ball of my
camera, the shutter clicks—the apotheosis is achieved! Like a
priestess, representative of her inventive sex, the graceful woman
stands: at her feet the hearth-fire she has given us beside her the
invention she has devised for us, in the background the home she has
built for us.
At Newala, also, I have had the manufacture of pottery carried on
in my presence. Technically the process is better than that already
described, for here we find the beginnings of the potter’s wheel,
which does not seem to exist in the plains; at least I have seen
nothing of the sort. The artist, a frightfully stupid Makua woman, did
not make a depression in the ground to receive the pot she was about
to shape, but used instead a large potsherd. Otherwise, she went to
work in much the same way as Salim’s mother, except that she saved
herself the trouble of walking round and round her work by squatting
at her ease and letting the pot and potsherd rotate round her; this is
surely the first step towards a machine. But it does not follow that
the pot was improved by the process. It is true that it was beautifully
rounded and presented a very creditable appearance when finished,
but the numerous large and small vessels which I have seen, and, in
part, collected, in the “less advanced” districts, are no less so. We
moderns imagine that instruments of precision are necessary to
produce excellent results. Go to the prehistoric collections of our
museums and look at the pots, urns and bowls of our ancestors in the
dim ages of the past, and you will at once perceive your error.
MAKING LONGITUDINAL CUT IN
BARK

DRAWING THE BARK OFF THE LOG

REMOVING THE OUTER BARK

 BEATING THE BARK

WORKING THE BARK-CLOTH AFTER BEATING, TO MAKE IT

SOFT

MANUFACTURE OF BARK-CLOTH AT NEWALA

To-day, nearly the whole population of German East Africa is
clothed in imported calico. This was not always the case; even now in
some parts of the north dressed skins are still the prevailing wear,
and in the north-western districts—east and north of Lake
Tanganyika—lies a zone where bark-cloth has not yet been
superseded. Probably not many generations have passed since such
bark fabrics and kilts of skins were the only clothing even in the
south. Even to-day, large quantities of this bright-red or drab
material are still to be found; but if we wish to see it, we must look in
the granaries and on the drying stages inside the native huts, where
it serves less ambitious uses as wrappings for those seeds and fruits
which require to be packed with special care. The salt produced at
Masasi, too, is packed for transport to a distance in large sheets of
bark-cloth. Wherever I found it in any degree possible, I studied the
process of making this cloth. The native requisitioned for the
purpose arrived, carrying a log between two and three yards long and
as thick as his thigh, and nothing else except a curiously-shaped
mallet and the usual long, sharp and pointed knife which all men and
boys wear in a belt at their backs without a sheath—horribile dictu!
[51]
Silently he squats down before me, and with two rapid cuts has
drawn a couple of circles round the log some two yards apart, and
slits the bark lengthwise between them with the point of his knife.
With evident care, he then scrapes off the outer rind all round the
log, so that in a quarter of an hour the inner red layer of the bark
shows up brightly-coloured between the two untouched ends. With
some trouble and much caution, he now loosens the bark at one end,
and opens the cylinder. He then stands up, takes hold of the free
edge with both hands, and turning it inside out, slowly but steadily
pulls it off in one piece. Now comes the troublesome work of
scraping all superfluous particles of outer bark from the outside of
the long, narrow piece of material, while the inner side is carefully
scrutinised for defective spots. At last it is ready for beating. Having
signalled to a friend, who immediately places a bowl of water beside
him, the artificer damps his sheet of bark all over, seizes his mallet,
lays one end of the stuff on the smoothest spot of the log, and
hammers away slowly but continuously. “Very simple!” I think to
myself. “Why, I could do that, too!”—but I am forced to change my
opinions a little later on; for the beating is quite an art, if the fabric is
not to be beaten to pieces. To prevent the breaking of the fibres, the
stuff is several times folded across, so as to interpose several
thicknesses between the mallet and the block. At last the required
state is reached, and the fundi seizes the sheet, still folded, by both
ends, and wrings it out, or calls an assistant to take one end while he
holds the other. The cloth produced in this way is not nearly so fine
and uniform in texture as the famous Uganda bark-cloth, but it is
quite soft, and, above all, cheap.
Now, too, I examine the mallet. My craftsman has been using the
simpler but better form of this implement, a conical block of some
hard wood, its base—the striking surface—being scored across and
across with more or less deeply-cut grooves, and the handle stuck
into a hole in the middle. The other and earlier form of mallet is
shaped in the same way, but the head is fastened by an ingenious
network of bark strips into the split bamboo serving as a handle. The
observation so often made, that ancient customs persist longest in
connection with religious ceremonies and in the life of children, here
finds confirmation. As we shall soon see, bark-cloth is still worn
during the unyago,[52] having been prepared with special solemn
ceremonies; and many a mother, if she has no other garment handy,
will still put her little one into a kilt of bark-cloth, which, after all,
looks better, besides being more in keeping with its African
surroundings, than the ridiculous bit of print from Ulaya.
MAKUA WOMEN