THE EFFECTS OF CROSS AND SELF FERTILISATION IN THE VEGETABLE KINGDOM, DARWIN

V THE EFFECTS CROSS AND SELF FERTILISATION VEGETABLE KINGDOM Bi CHAELES DABWIN, M.A., F.E.S., ETC D NEW YORK: APPLETON AND COMPANY, 1, 3, and BOND STEEET 1892 TS\\e> — CONTENTS CHAPTER I Introductory Remarks Various means which favour or determine the cross-fertilisation of plants — Benefits derived from cross- fertilisation tion favourable to the propagation of-the species — — Self- fertilisa- Brief history — Object of the experiments, and the manner in which they were — value of the measurements The experiments on during successive generations — Nature of the relationship of the plants in the generations — Uniformity of the conditions which the plants — were subjected — Some apparent and some causes of Amount of pollen employed — Arrangement of the work — of the subject tried Statistical carried several later to real Importance of the conclusions error Page 1-27 CHAPTER II CoNVOLVtrLACE^S Fpomcea purpurea, comparison of the height and fertility of the crossed and self-fertilised plants during ten successive generations Greater constitutional vigour of the crossed plants The effects on the offspring of crossing different flowers on the — — same instead of plant, effects of self-fertilised vigour, crossing distinct a cross with a fresh stock and generations plant fertility named Hero individuals — The descendants — Summary — The of the on the growth, of the successive crossed and self-fertilised — Small amount of pollen in the anthers of the self- CONTENTS IV the later generations, and fertilised plants of their first-produced flowers the sterility of — Uniform colour the — The advantage of flowers from-' a produced by the self-fertilised plants cross between two distinct plants depends on their differing in Pa^e 28-62 constitution CHAPTEE IIL ScrophulariacejE, Geskeeiace^:, Labiate, etc Mimulus luteus self-fertilised of a new, cross plants of the tall, between fresh stock and height, vigour, ; — and highly first fertility of the crossed four generations self-fertile variety self-fertilised plants — and — Appearance — Offspring from a Effects of a cross with Effects of crossing flowers on the same plant a — luteus — purpurea, — on the same of — Linaria — Verbascum thapsus — plant — Vandellia nummularifolia — Cleistogene — Gesneria pendulina — Salvia coccinea — Origanum — Thunbergia the plants by 63-97 Summary on Mimulus of the crossed plants Digitalis superiority crossing flowers Effects vulgaris Calceolaria flowers vulgare, great increase of crossed alata stolons CHAPTEE IV Crucifer^, Papaverace.e, Resedacej3, etc Biassica oleracea, crossed and self-fertilised plants — Great a cross with a fresh stock on the weight of the offspring — — effect of — Iberis umbellata Papaver vagum Eschscholtzia californica, seedlings from a cross with a fresh stock not more vigorous, but more fertile than the self-fertilised seedlings Eeseda lutea and — odorata, tricolor, many individuals sterile with their wonderful effects — of a cross own pollen — Adonis — Viola — aestivalis Delphinium consolida Viscaria oculata, crossed plants hardly taller, but more fertile than the self-fertilised Dianthus caryophyllus, crossed and self- fertilised plants compared for four generations Great effects of a cross with a fresh stock Uniform colour of the flowers on the self-fertilised plants Hibiscus africanus 98-141 — — — — V CONTENTS CHAPTEE V Geraniace^e, Legtjmtnos.e, Onagrace^;, etc Pelargonium zonale, a cross between plants propagated by cuttings Limnanthes douglasii Tropseolum minus does no good Phaseolus multiflorus and vulLupinus luteus and pilosus garis Lathyrus odoratas, varieties of, never naturally inter- — — England cross in — Pisum — — — sativum, varieties of, — rarely inter- but a cross between tbem highly beneficial Sarothamnus Ononis minutissima scoparius, wonderful effects of a cross Summary on the Leguminosa? cleistogene flowers of Clarkia cross, — — — Bartonia aurea — Passi elegans — — — Apium petrosesativa — Specularia flora gracilis — — Lactuca linum Scabiosa atropurpurea speculum Lobelia ramosa, advantages of a cross during two generations Lobelia fulgens Nemophila insignis, great advantages of a cross Borago officinalis Nolana prostrata Page 142-187 — — — CHAPTEE — VI JSolanace^:, Primulaceje, Polygone^:, eto Petunia violacea, crossed and self-fertilised plants compared for four — with a of a stock — Uniform colour of the flowers on the plants of the fourth generation — Nicotiana tabacum, crossed and generations Effects fresh cross self-fertilised plants of equal height — Great self-fertilised effects of a cross with a distinct sub- variety on the height, but not on the spring to — Cyclamen fertility, of the off- persicum, crossed seedlings greatly superior Anagallis collina sell-fertilised Primula veris the — — — Equal-styled variety of Primula veris, fertility of, greatly stock —Eagopyrum esculentum with a — Canna warscewiczi, crossed and plants of equal height — Zea mays — Phalaris canadensis increased — Beta by a cross vulgaris fresh self-fertilised 188-237 CONTENTS VI CHAPTER VIL Summary of the Heights and Weights of the Crossed and Self-fertilised Plants Number of species and plants measured — Tables — given — Pre- liminary remarks on the offspring of plants crossed by a fresh, Thirteen cases specially considered The effects of stock — by another crossing a self-fertilised plant either plant or by an self-fertilised intercrossed plant of the old stock — Summary — Preliminary remarks on the crossed and the same stock — The twenty-six exceptional which the crossed plants did not exceed — Most these shown height the greatly not exceptions the that be — Relative weights of the — Summary of the results self- fertilised plants of cases considered, in in to to beneficial crossed of and cases of self-fertilised real rule cross-fertilisation is results Page 238-284 self-fertilised plants CHAPTER VIII Difference between Crossed and Self-fertilised Plants in Constitutional Vigour and in other respects — The — — Competition with other kinds more premature death — Crossed plants — Negative of the on the same plant — Cases described — generations Transmission the good a — parentage — Greater constitutional vigour of crossed plants crowding tilised plants Self-fer- liable to generally flower before intercrossing effects self- fertilised flowers of Effects effects of great of plants effects of of crossing plants of cross to later closely related Uniform colour of the flowers on plants self-fertilised during several generations and cultivated under similar conditions 285-31J CHAPTER The Effects of IX Cross-fertilisation and Self-fertilisation on the Production of Seeds and self-fertilised parentage, both lots same manner Fertility of the parentcrossed and self-fertilised, and of their crossed Fertility of plants of crossed being fertilised in the plants when first — CONTENTS Vll and self-fertilised offspring when again crossed and self-fertilised Comparison of the fertility of flowers fertilised with their own pollen and with that from other flowers on the same plant — — Self-sterile plants — Causes of — of highly self-fertile varieties some self-sterility — The appearance Self-fertilisation apparently in respects "beneficial, independently of the assured produc- tion of seeds — Relative weights and rates of germination of Page 312-355 seeds from crossed and self-fertilised flowers CHAPTEE Means of X Fertilisation and fertility of plants when insects means by which flowers are cross-fertilised Sterility excluded — The — Structures favourare — Eelation between the structure and — The means by which flowers tages of with pollen from a plant — Greater — Conversion power such pollen — Anemophilous anemophilous entomophilous — Origin of nectar — Anemophilous plants generally have sexes separated — — Trees Conversion diclinous hermaphrodite able to self-fertilisation conspicuousness of flowers, the visits of insects, and the advancross-fertilisation are distinct fertilised of fertilising species of species into their of into flowers 356-414 often have their sexes separated CHAPTER XL The Habits of Insects in relation to the Fertilisation of Flowers Insects visit the flowers of the Cause of this flowers of the same species as long as they can — Means by which — Sudden same habit species bees recognise secretion of nectar — — the — Nectar of certain flowers unattractive to certain insects Industry of bees, and the number of flowers visited within a short Perforation of the corolla by bees Skill shown in the time Hive-bees profit by the holes made by humble-bees operation The motive for perforating flowers to save Effects of habit Flowers growing in crowded masses chiefly perforated time — — — — — — 415-435 CONTENTS VU1 CHAPTEE XII Gekeeal Eesults Cross-fertilisation proved to jurious — Allied cross-fertilisation is The benefits and be beneficial, and self-fertilisation in- species differ greatly in the means by which favoured and self-fertilisation avoided evils of the two — processes depend on the degree of differentiation in the sexual elements — The evil effects not — due to the combination of morbid tendencies in the parents Nature of the conditions to which plants are subjected when growing near together in a state of nature or under culture, and Theoretical considerations with the effects of such conditions — respect to the interaction of differentiated sexual elements Practical lessons — Genesis of the two sexes — — Close correspon- dence between the effects of cross-fertilisation and self-fertilisation, and of the legitimate and illegitimate unions of heterostyled plants, in comparison with hybrid unions Page 436-469 Indbx Page 471—482 THE EFFECTS OF CEOSS AND SELF-FEKTILISATION CHAPTER I INTRODUCTORY KeHARKS Various means which favour or determine the cross -fertilisation of plants Benefits derived from cross-fertilisation Self-fertilisation favourable to the propagation of the species Brief history of the subject Object of the experiments, and the manner in which they were tried Statistical value of the measurements The experiments carried on during several successive generations Nature of the relationship of the plants in the later generations Uniformity of the conditions to which the plants were subjected Some apparent and some real causes of error Amount of pollen employed Arrangement of the work Importance of the conclusions — — — Theee is flowers of — — — — — — — — — weighty and abundant evidence that the most kinds of plants are constructed so as be occasionally or habitually cross-fertilised by pollen from another flower, produced either by the same plant, or generally, as we shall hereafter see to reason to believe, by a distinct plant Cross-fertilisation is sometimes ensured by the sexes being separated, and in a large number of cases by the pollen and stigma of the same flower being matured at different Such plants are called dichogamous, and have times been divided into two sub-classes proterandrous : species, ; INTRODUCTORY REMARKS Chap I is mature before the stigma, and proterogynous species, in which the reverse occurs this latter form of dichogamy not being nearly so in which the pollen common as the other sured, in many cases, Cross-fertilisation is also en- by mechanical contrivances of wonderful beauty, preventing the impregnation of the There is a small class of flowers by their own pollen which I have called dimorphic and trimorphic, but to which Hildebrand has given the more appro- plants, name priate of heterostyled ; this consists of class plants presenting two or three distinct forms, adapted for reciprocal fertilisation, so that, like separate sexes, they can hardly fail plants with to be intercrossed The male and female organs of some flowers are irritable, and the insects which touch them get dusted with pollen, which is thus transported Again, there is a class, in which the to other flowers ovules absolutely refuse to be fertilised by pollen from the same plant, but can be fertilised by pollen from any other individual of the same species There are also very many species which are partially sterile with in each generation their own pollen Lastly, there is a large class in which the flowers present no apparent obstacle of any kind to self-fertilisation, nevertheless these plants are frequently intercrossed, owing to the prepotency of pollen from another individual or variety over the plant's As own pollen plants are adapted by such and effecmight have been inferred from this fact alone that they derived some great advantage from the process and it is the object of the present work to show the nature and importance of the benefits thus derived There are, however, some tive means diversified for cross-fertilisation, it ; exceptions to the rule of plants being constructed so as to allow of or to favour cross-fertilisation, for some ;; INDEX 478 MELASTOMACEiE Melastomacex, 298 affi- 456 Mimulus luteus, effects of crossing, crossed and self-fertilised 10; plants, 64-70 measurements, 7078 cross with adistinct stock 72intercrossed on same plant 75 ; ; ; 75-78 78-81 summary of observations of experiments, 259-261 superiority of crossed plants, 286 simultaneous flowering, 294, 296 of intercrossing, 301 effects uniform colour of self-fertilised beeds, 315, 319, 322, 324 307; highly self-fertile, 348, 369 ; prepotency of other pollen, 392, 397 ; ; roseus, 63 Miner, Mr., red clover never sucked by hive-bees in the United States, 361 Mirobilis, dwarfed plants raised by using too few pollen-grains, 298 number of grains necessary for fertilisation, 377 ; Mitchell, Dr., on first-cousins inter- marrying, 460 Monochoetum ensiferum, 364 Moure, Mr., on Cinerarias, 335 Fritz, on Posoqueria Midler, fragrans, 5, 391 experiments on hybrid Abutilons and Bignonias, ; of large number 306; orchidaceous genera sterile in Bignonia their native home, also 305, Tabernxmontana and echinata, of Eschscholtzia Abutilon californica, 332, 342 darwinii, 334; experiments in 331 ; sterility ; self-fertilisation, plants, 341 340; ; ; ; lowland and alpine species, 375; structure of plants adapted to cross and self- fertilisation, 380; large conspicuous flowers more frequently visited by insects than small inconspicuous ones, 383; Solatium generally Unattractive Lamium album, to insects, 387 on anemophilous plants, 389 400 fertilisation of Plantago, secretion of nectar, 405 401 instinct of bees sucking nectar, 415; bees frequenting flowers of the same species, 416; cause of it, 419; powers of vision and discrimination of bees, 422 Midler, Dr H„ hive-bees occasionally perforate the flower of Erica tetralix, 427; calyx and corolla of Ehinantlius alecterolophus bored by Bombus mastrucatus, 432 ; ; ; ; Munro, Mr., some species of Onciclium and Maxillaria sterile with own pollen, 331 Myrtacex, 412 selt'-sterile N by means ; ; cross-fertilisation imperfectly deveof birds, 371 loped male and female Termites, food-bodies in Cecropia, 404 3S0 idler, Hermann, fertilisation of flowers by insects, on 6, Digitalis purpurea, 82; Calceo; ; M ; N'ageli ; ; flowers of Pisum sativum, 160, 161; on Sarothamnus scaparius not secreting nectar, 164 Apium petroselinum, 172 Borago officinalis, 1S5; red clover visiled by hive-bees in Germany, 361 insects rarely visiting Fumaria comparison of officinalis, 366 147 incapacity of pollentubes to penetrate the stigma, 342 ; ; Mercurialis annua, 413 Miller, Professor, on chemical nity, Linaria vulgarity, 88 87 Verbascum nigrum, 89 the common cabbage, 98; Pavacer dubium, 107; Viola tricolor, 123, 124; structure of Delphinium coiisoliila, 129; of Lupinns lutea, laria, Melilotus officinalis, 360 ; on odours attracting insects, 374 sexual relations, 09 Natural selection, effect upon selfsterility and self-fertilisation, 345, 346 Naudin on number of pollen-grains necessary for fertilisation, 24 Petunia violaeea, 188 ; INDEX 479 PHASEOLUS Nectar regarded as an excretion, 402 Nenwphila insignis, 182; measurements, 1S3-1H5; early flowering of crossed plant, 293 effects of cross and self-fertilisation, 303; seeds 316,323 Nepeta glechoma, 41 Nicotiana glutinosa, 210 tabacum, 203 measurements, 205-208; cross with fresh stock, 210; measurements, 212-215; : ; ' summary of experiments, 266, 267, 279 superiority of crossed plants, 288-290; early flowering, 293295; seem intercross, 160; summary of experiments, 264, 162; 278; self-fertile, 367 greater Plants, crossed, tutional vigour, 285 ing of crossed, 293; seeds, 317; prepotency of self-fertility, 351 dark red polyanthus, 396 Primulacex, 215 Pringlea, 408 Proteacex of Australia, 412 Prunus avium, 403 ; laurocerasus, 401 Pteris aquilina, 404 consti- Fleroma, 364 Polemoniacex, 182 E Pollen, relative fertility of flowers crossed from a distinct plant, or difference with their own, 320 of results in Nolana prostrata, 321, 323 ; crossed and self-fertilised again crossed from a plants, distinct plant and their own polsterile with their own, len, 324 ; ; 338number of 376 grains in Dandelion, Pseony, and Wistaria sinensis, 376 number necessary for fertilisation, 377 transported from flower to flower, 377 prepotency, 391-400 abori330-338 loss 340 ; ; seuii-self-sterile, of, ; ; ; ; ginally the sole attraction to insects, 402 ; quantity produced by anemophilous plants, 405 Polvanthus prepotency over cowslip, 396 Polygoneie, 228 Posoqneria fiagrans, 5, 391 Potato, 387 Poterium sanguisorba, 407 Potts, heads of Anthornis melanura covered with pollen, 371 Primrose, Chinese, 225 Primula elatior, 422, 427 grandijlora, 378 mollis, 368 officinalis, 378 scotica 362 measuresinensis, 225, 279 early flowering of ments, 227 crossed, 293, 296 — veris (var officinalis'), 219 measurements, 221 result of experiments, 267, 268 early flower- — : Radish, 393 Ranunculaceie, 128 Ranunculus acris, 365 Raphanus sativus, 365, 393 Reinke, nectar-secieting glands of Prunus avium, 403 Eeseda lutea, 117; measurements, 118, 119; result of experiments, 339; self fertile, 365 odorata, 119; measurements, 120-123 self-fertilise J scarcely exceeded by crossed, 289 seeds, 316; want of correspondence between seeds and vigour of offspring, 328; result of experiments, 336; sterile and self-fertile, 358, 365 Resedaceas, 117 Rheum rhaponticum, 401 Rhexia glandulosa, 364 Rhododendron, spontaneous crossing, 394 ; ; Rhododendron azaloides, 432 Rhubarb 391, 401 Ribes aureum, 432 Riley, Mr., pollen carried by wind, 405; Yucca moth, 418 Rodders, Mr., secretion of nectar in Vandla, 408 Rye, experiment on pollen cf, 376 S , ; ; — ; ; Salvia coccmea, 93 measurements, early flowering of crossed, 93 ; ; 292; seeds, 315, 363 glutinosa, 427 self-sterile, 322; partially ;; 481 INDEX Salvia graliami, 420, 42S, 433 tenon, 302 Sarothamnus scoparius, 163; measure! 1- its, l-i 55-1 67 superiority of crossed seedli.igs, 28.i, 289 seed s 323: self-sterile, 3ti0 Seabiosa atro-purpurea, 172; measurements, 172, 173 Scarh t-runner, 150 1 ; Papaver somniferum, 108 Verbascum, 330 Onon cidium and Maxillaria, 331 Primula scoti a and Cortusa mat- Scott, J., ; steril.ty of ; ; 362 thioli, tum napellus, 428 importance of insects in fertilising flowers, 455 Stachi/s coccinea 426, 428, 433 Stellaria media, 367 Strachey, General, perforated flowers in ttie Hima aya, 434 ; Strawberry, 3.*4 by the Nectarini371 Structure of plants adapted to cross and self fertilisation, 380 Swale, Mr., garden lupine not visited by bees in New Zealand, 150 Sweet-pta, 153 Strelitzia fertilised deie, Scrophulariacete, 63 appearance Self-feitile varieties, of, 347-351 mechanical struccheck, 382 Self-fertilisation, ture to ecliinata 331, 362 asuretnents of heighrs, weights, and fertility of plants, Tabernannontana wide Table.- of throughout the differvegetable kingdom, 341 cause of ence in plants, 342 Termites, Self-sterile plants, f.29-347; dlstribut on ; ; 343 self-sterility, ; by affected changed 344-346 conditions, necessity of differentiation in the sexual elements 317 Senecio emeritus, 335, 364 >5 maderensis, 335 populifoius, 335 heritieri, tussilaginis, 3:->5 Sliaipe, Mes>rs., precautions against 395 intei crossing, Snow-Hake, 176 Solanaceie, 1S8 Solanum tuberosum, 362, 387 Specularia per/oliata, 174 speculum, 174; measurements, 175, 176; crossed and self-fertiearly flowering of lised 216; crossed, 293 fertile, ; seeds, 323 ; self- 369 Spencer, Herbert, chemical affinity, Hi' 240-270 imperfectly developed males and female.s, 3S0 Thunbergia alata, 90, 277, 331 Tin me, 419 Tiuziiiami, on Solarium tuberosum, 362, 388 Tobacco, 203 Transmission of the good effects of a cro.-s to Jater generations, 303 Trees separated seXi-s, 411 Trifoliuui aroense, 367, 384 incaruatum, 361 minus, 368 pratense, 361 426, 434 prociinibens, 368 repens, 361 Tropxolum minus, 144 ; measure- ment-, 145 ; early flowering of 293 seeds, 316, 323 seeds, 323 tricolor, 427 Tulips, 394 Typha, 405 cro.ss d, ; ; 457 Spiranthes autumnalis, 391, 421 K, filiation of Viola 5, colours in flowers tricolor, 123 attract and guide insects, 372, 373 ; on Aristolochia, ill ; Aconi- Spreiiirel, C flowers by fei insects, ; ; Umbelliferx, 172 Urban, Ig., fertilisation of lupulina, 368 Medioago ; ; INDEX 482 VANDELLIA ZEA crossed and self-fertilised, 276 simultaneous flowering, 295; seeds, 316, 323; self-fertile, 367 V Yandellia nummularifolia, 90, 278 seeds, 315, H22 self-fertile, 369 ; Yauilla, secretion of nectar, 403 Verbascum hjehnitis, 89, 311, 309 nigrum, 89 3:;0, 3tl phceniceum, 330, 341, 364 thapsus, measurements, 89 90; self-fertile, 311,3 ,9 Vcilot on Convolvulus tricolor, 55; intercrossing of Nemophila, 183; of Leptosiphon, 394 Veronica agrestis, 369 chamvkdrys, 369 Uedernefolia, 369 Viciafaba, 360, 404 hirsuta, 367 : sa^'ua, 367, 403, 404 Victoria regia, 365 Villarsia parnassifolia, 390 Vilmorin on transmitting character to offspring, 451 Vinca major, r>62 rosea, 362 Viola canina, 357 tricolor, 123; measurements, 126, 127 ; superiority of crossed plants, period of 286, 289 flowering, 292, 296; effects of seeds, cross-fertilisation, 304 partially sterile, 358 ; 316, 325 corolla removed, 420 W Wallace, Mr., the beaks and faces of brush-tongned lories covered with pollen, 371 Wasps attracted by Epipactis latifolia, 375 Weights, relative, of crossed and self-fertilised plants, 244 283; and period of germination of seeds, 352-355 Wither, Mr., fertilisation of flowers with their own pollen, 341 WiLon, A J., superior vigour of crossed seedlings in Brassica campestris ruta baga, 353 Wistaria sinensis, 376, 427 Y Yucca moth, 448 Z : ; ; Violacex, 123 Viscaria oculata, ment, 132 130; measureaverage height of measurements, 16, 233 16-18, 234; difference of height between crossed and self-fertilise d, 288; eariy flowering of crossed, 293 self-fertile, 369 prepotency of other pollen, 397 Zea mays, ; ; ; D APPLETON & CO.'S PUBLICATIONS JOHN TYNDALL'S WORKS ESSAYS ON THE FLOATING MATTER OF THE AIR, in Relation to Putrefaction and ON FORMS OF 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