FRUITS OF ANGIOSPERMS
by
Prof. Dr. I N G R I D R O T H
Universidad Central de Venezuela, Caracas
With 232 figures
1977
GEBRUDER BORNTRAEGER
• BERLIN • STUTTGART
Contents
A. General part
1
1.1
1.2
2
3
4
4.1
5
6
6.1
6.2
6.3
6.4
6.5
7
8
9
10
11
12
13
The concept "fruit"
Definition of the fruit
Classification of fruits
The carpel
Fruit symmetry
The pericarp
Definition of exo-, meso-, endocarp
Relation between anatomical structure and functions of the pericarp .
.
Fruit growth
Growth stimulation
Cell division
Cell enlargement
Periods of growth
Cell size in relation to fruit size
Meristems and fruit growth
Cell and tissue differentiation
Cell contents
Maturation
Senescence
Texture and edibility of fruits
Pattern formation
Outer epidermis 37, Inner epidermis 37, Interwoven pattern 38, Formation of radial cell rows 38, Starlike arrangement of cells 38, Giant
cells 38
1
1
1
1
6
8
9
9
11
12
13
14
15
16
19
21
26
27
30
33
34
37
B. Tissues composing the pericarp
39
14
Epidermis
14.1 Outer epidermis
Shape of epidermal cells
Cell wall structure
Cutine and wax formations
Cell contents
Multiseriate outer epidermis
39
39
41
41
42
42
42
X
14.2
14.3
14.4
14.5
15
16
17
17.1
17.2
17.3
17.4
18
19
19.1
19.2
20
21
21.1
21.2
Contents
Inner epidermis
Multiseriate inner epidermis
Differentiation of the endocarp
Stomata
Occurrence of stomata
Stomata distribution and density
Stomata development
Anomalous stoma formation and reduction of stomata
. . . .
Function of stomata
Trichomes
Shape of trichomes (unicellular hairs, pluricellular hairs) . . . .
Internal or locular hairs
Development of hairs
Density of hairs
Systematic value of hairs
Function of hairs
Emergences
Periderm and lenticel formation
Hypodermis
Parenchyma
Special cell arrangements and cell patterns
Aerenchyma
Air containing parenchyma
Cell contents
Collenchyma
Sclerenchyma
Sclereids
Cell shape
Distribution of sclereids
Special stone cell patterns in Solanaceae
Taxonomic and phylogenetic value of stone cell concretions . . .
Transitional forms between sclereids and fibers or parenchyma cells .
.
Fibers
Shape of fibers
Distribution and arrangement of fibers
Origin and differentiation of stone cells and fibers
Vascular system
Secretory structures
External secretory structures
Nectaries hydathodes, glandular hairs
Internal secretory structures
Secretory cells 94, Secretory cavities 95, Secretory canals 96, Laticifers 97
44
47
48
49
51
52
54
54
57
57
57
60
62
62
63
63
65
66
69
70
71
73
73
74
74
75
77
77
79
80
80
82
82
83
83
86
87
93
93
93
94
C. Structures beyond the pericarp
98
22
22.1
98
98
Placentae and septs
Placentae and septs as sources of pulp formation
Contents
XI
22.2
False partition walls and seed pockets
23
The inferior ovary
100
24
The fruits of cacti
106
25
25.1
25.2
25.3
Accessory fruit parts
Sterile projections of the gymnoecium ("Grannen")
Receptacle
Sepals
Pappus
Petals
Anthocarps
Stamens
Fruit scales
Prophylls
Pedicel and peduncle of the fruit
Parts of the inflorescence
Inflorescence axis 142, Glumes spikclets 142, Subtending bracts 146,
Involucral bracts of inflorescences 146
118
119
121
122
126
128
128
131
131
131
132
142
25.4
25.5
25.6
25.7
25.8
25.9
100
D. Fruit dehiscence
148
26
26.1
26.2
26.3
26.4
26.5
26.6
26.7
148
148
152
153
153
156
163
166
166
27
27.1
27.2
28
Types of dehiscence
General considerations
Ventricidal ("marginicidal") dehiscence
Septicidal dehiscence
Loculicidal dehiscence
Dehiscence of the legume
Pore capsule
Circumscissile dehiscence
Pyxidia which develop sclerenchyma and a zone of mechanical weakness
Pyxidia which dehisce by formation of meristematic cells in combination with lignified cells
168
Mechanisms of dehiscence
Hygroscopic mechanisms
Xerochasy
Hygrochasy
Turgor mechanisms
Spluttermechanisms (Spritzmechanismen)
Throwing mechanisms of turgcscent fleshy fruits (Schleudcrmechanismen)
Lever catapults (Hebelschleudern)
Cyclanthcra cxplodens 182, Impaticns 1S2. Cardammc 1S4, Corydalis 184, Lathracu 184
Squeezing catapults (Quctschschleudcrn)
Dorstenia 185, Arccuthobium 185
Pushing catapults (StofSschleudern)
Polygonum 187
171
171
172
175
181
1S1
182
1S2
Indehiscent fruits with delayed dehiscence
187
185
187
XII
Contents
E. Types of fruits
191
29
The dehiscent fruit: the capsule sensu lato
29.1 General considerations
29.2 Special examples
Solanaceae
Caryophyllaceae
The legume
Vascular anatomy
Development
Formation of the "seed-cushion':
Formation of false septs
Follicular types with ventricidal dehiscence only
Evolutionary trends in fruit development of Leguminosae
Cruciferae (Silique and silicle)
Acanthaceae
Scrophulariaceae
Orchidaceae
Liliaceae
Linum usitatissimum
Fleshy capsules
Myristica fragrans (the nutmeg)
Punica granatum (the pomegranate)
Scyphostegia borneensis
191
191
194
197
200
203
209
210
210
212
215
216
218
225
228
231
231
233
235
235
239
244
.
30
The indehiscent sclerocarpium: the nut
30.1 The nut sensu stricto
30.2 The achene
The genus Centcturea
Distribution of the parenchyma in the pericarp
The aerenchyma
The sclerenchyma
The "carbon layer" (,,Kohleschicht") or black pigment layer .
The hypodermis
Secretory structures
Cell contents
Formation of intercellular spaces
The epidermis
Slime cells
Slime trichomes
Twin hairs and other trichomes
Periderm
Emergences
Vascular system
The apical plate
The receptacle of the inflorescence
The integument
Structure of the pappus
Chaffy bracts and involucral leaves
.
.
.
.
244
246
258
261
264
266
267
270
271
273
273
274
274
276
277
278
280
281
281
281
283
284
284
286
Contents
30.3
30.4
30.5
30.6
30.7
30.8
31
31.1
31.2
31.3
31.4
31.5
31.6
31.7
31.8
31.9
31.10
31.11
31.12
31.13
XIII
The abscission region
Heterocarpous inflorescences
Taxonomic significance of the pericarp structure
The caryopsis of Gramineae
General structure
2,ea mays
Pericarp development
Semipermeable membrane
Bracts (glumes and palets) taking part in fruit formation . . . .
The schizocarpium of Umbelliferae
General pericarp structure
The schizocarpium of Hydrocotyloideae
Secretory spaces
Interpretation of the inferior ovary
The carpophore
Classification based on fruit anatomy
Evolutionary trends of the schizocarp and derivation from fruits of
Araliaceae
The mericarps or nutlets ("Klausen") of Boraginaceae
The mericarps or nutlets ("Klausen") of Labiatae
Anacardium occidental
Winged fruits (samaras)
287
290
290
291
291
293
296
304
307
311
313
314
318
32C
320
324
The fleshy-stony indehiscent fruit: the drupe (sarco-sclerocarpium) .
.
General characteristics
Zonation of meso- and endocarp
The Prunus fruits
Prunus domestica (The plum)
Prunus cerasus (The sour cherry)
Prunus persica (The peach)
Vascular system
Cell contents
Tissue texture
Size and form of fruits
Growth periods
Species of Rubus
Rubus idaeus (Red raspberry)
Rubus jruticosus (Black raspberry)
Rubus strigosus (American red raspberry)
Olea europaea (The olive)
Coffea arabica, C. canephora (The coffee)
Cocos nucifera (The coconut)
Other palm fruits
Casimiroa edulis (The white sapote)
Mangijera indica (The mango)
Ribes (Saxifragaceae)
Fruits of Ericaceae: Vaccinium, Gaylussacia
Symphoricarpus nvulans (cit. 5. racemosus). Conns mas, Gcum urbanum
368
368
372
325
330
336
344
358
374
374
376
378
"
379
u
3
^
3SI
381
3S3
383
3S6
38S
uU
4
|;
A2Q
4 3
~
*~424
XIV
Contents
31.14 Juglandaceae
Carya illinoensis (cit. Hicoria pecan) 425, Carya ovata and Carya
cordiformis 427, Juglans 428
425
32
The fleshy indehiscent fruit: the berry (sarcocarpium)
32.1 General characteristic
32.2 Types of berries
Ribes
Phoenix dactylifera (The date)
Acbras zapota (The sapote)
Persea americana (The avocado)
Solanaceae
General characteristics
Lycopersicon esculentum (The tomato)
Capsicum annuum (The red Pepper)
Musa acuminata and M. balbisiana — cultivars (The banana) . . .
Fruits of Cucurbitaceae (Type: Pepo)
Exocarp
Cork formation
Mesocarp
Endocarp
Vascular bundles
Carica papaya (The paw paw)
Anatomy of the ripe fruit
Development of the fruit
Theobroma cacao (The Cacao)
Development of the fruit
Anatomy of the mature fruit
Species of Citrus (Type: hesperidium)
General structure of the Citrus fruit
Development of the Citrus fruit
Cell contents
Other members of Rutaceae
Loranthaceae
The clove (dried flower buds of Syzygium aromaticum or Eugenia
caryophyllata
431
431
433
433
434
435
445
451
451
454
457
461
471
472
475
475
477
477
477
480
480
483
483
489
494
494
502
511
516
517
33
Aggregate fruits and apocarpous gynoecia
33.1 Ranunculaceae
33.2 Annonnaceae
Annona cherimola (The cherimoya)
Morphology
Anatomy
Outer epidermis 526, Mesocarp 526, Inner epidermis 526, Vascular
anatomy 526, Cell contents 527
33.3 Rosaceae
Fragaria sp. (The strawberry)
Rosa
522
522
524
524
524
526
519
527
530
531
Contents
XV
533
533
533
540
540
33.4
The pome fruits
Malus sylvestris (The apple)
Vascular pattern of the apple flower
Histogenesis
Histology of the ripe apples
Epidermis 540, Cork and lenticels 541, Hypodermis 543, Fruit
flesh 543, Endocarp 543
Calyx
Texture of flesh and air spaces
Cell contents
Growth rate, cell size and cell number
Fruit abscission
Pyrus communis (The pear)
Vascular supply
Histogenesis
Histology
Outer epidermis 548, Cork formation 548, Hypodermis 550,
Flesh 550, Scbreid differentiation 550, Endocarp 550
Growth rate
The pseudo-apocarpous fruit of Ochna
34
34.1
34.2
34.3
Infrutescences (collective or multiple fruits)
Ananas comosus (The pineapple)
Artocarpus altilis (the bread fruit) and related species
Ficus cariaca and related species (The fig)
543
543
543
544
546
546
546
546
548
552
552
553
553
557
560
F. Fruit abscission
565
G. Special dispersal structures and heterodiaspory
572
35
Elaiosomes
572
36
36.1
36.2
36.3
36.4
36.5
Floating fruits
Floating tissue without air spaces
Floating tissue in the form of aerenchyma
Floating bladders filled with air
Floating tissue developed outside the pericarp
Protective layers of floating fruits
579
579
581
582
582
582
37
37.1
37.2
37.3
Pneumatocarpia and flying organs
Pneumatocarpia
Flying organs with airspaces
Hairs serving as flying organs
583
583
585
585
38
Dispersal organs in the form of hooks or burs
587
39
39.1
Heterodiaspory
Heterocarpy
Heterocarpous Compositae
Heterocarpy in other families
587
591
594
597
XVI
39.2
39.3
39.4
39.5
Contents
Amphicarpy
Heterocarpidy
Heteromericarpy
Seed pockets
597
598
598
599
H. Taxonomic use of the pericarp structure
600
I. Phylogeny of the fruit
603
General view
Special examples
K. Fruits of cultivated plants and polyploidy
Bibliography
Index of authors
Index of latin plant and animal names
General index
603
605
609
615
643
649
666