Chapter 15

I. RHIZOFLAGELLATA (PANTOSTOMATA)Food taken in by pseudopodia at any part of the body.Order 1.—HOLOMASTIGACEAE. Body homaxial with uniform flagella.Multicilia(Cienkowski);Grassia(Fisch, in frog’s blood and gastric mucus).Order 2.—RHIZOMASTIGACEAE. Flagellum 1, 2 or few, diverging from anterior end.Mastigamoeba(F.E. Schulze).II. EUFLAGELLATAFood taken in at one or more definite mouth-spots, or by a true mouth, or by absorption; or nutrition holophytic.Order 1.—PROTOMASTIGACEAE. Contractile vacuole simple, one or more, or absent; either holozoic, ingesting food by a mouth-spot (or 2 or more), saprophytic, or parasitic.Family 1.—Oicomonadidae. Flagellum 1, sometimes with a tail-like posterior prominence passing into a temporary flagellum, but without other cytoplasmic processes.Oicomonas(Kent);Cercomonas(Dujardin) (Fig. 1,32, 33);Codonoeca(James-Clark), with a gelatinous theca.Family 2.—Bicoecidae. Differs fromOicomonadidaein a unilateral proboscidiform process next the flagellum; often thecate and stalked, forming branched colonies, like Choanoflagellates in habit.Bicoeca(J.-Cl.),Poteriodendron.Family 3.—Choanoflagellidae(Choanoflagellata, Kent; Craspedomonadina, Stein). As in previous families, but with flagellum surrounded by an obconical or cylindrical rim of cytoplasm, at the base of which is the ingestive area. The cells of this group have the morphology of the flagellate cells (choanocytes) of sponges. They are often colonial, and in the gelatinous colony ofProterospongia, the more internal cells (Fig. 2,15) pass into a definite “reproductive state.” Many stalked forms are epizoic on Entomostracan Crustacea.(a) Naked forms often stalked:Monosiga(Kent), stalked solitary;Codosiga(Kent) (Fig. 2,3), stalked social;Desmarella(Kent), unstalked, andAstrosiga(Kent), stalked, form floating colonies.(b) Forms enclosed in a vase-like shell:Salpingoeca(J.-Cl.); (Fig. 2,1, 6, 7) recalling the habit ofMonosigaandCod siga;Polyoecaforming a branched free swimming colony.(c) Forms surrounded by a gelatinous sheath:Proterospongia(Kent) (Fig. 2,15);Phalansterium(Cienk.) (Fig. 1,12), has a slender cylindrical collar, and a branching tubular stalk.Family 4.—Haemoflagellidae. Forms with a complex nuclear apparatus, and a muscular undulating membrane with which one or two flagella are connected, parasitic in Metazoa (often in the blood).Trypanosoma(Gruby) (Fig. 1,21, 22),Herpetomonas(Kent),Treponema(Vuillemin)(=Spirochaete, auctt., nec. Ehrbg.).Family 5.—Amphimonadidae. Flagella 2 anterior, both directed forward, equal and similar; in stalk sheath, &c., often recalling Choanoflagellata,Amphimonas(Kent),Diplomitus(Kent);Spongomonas(St.), with thick branching gelatinous sheath.Family 6.—Monadidae. Flagella 2 (3), anterior all directed forwards, one long the other (or 2) accessory, short.Monas(St.);Anthophysa(Bory) (Fig. 2,12, 13), with the stalk composed of the accumulation of faeces at the hinder end of the cells of the colony.Family 7.—Bodonidae. Flagella 2 (or 3) 1 anterior, the other (1 or 2) antero-lateral and trailing or becoming fixed at the end to form a temporary anchor.Bodo(Ehrb.) (figs. 1,23-26and 2,10).B. lensis the “hooked” andB. saltansthe “springing monad” of Dallinger and Drysdale;Dallingeria(Kent) with a pair of antero-lateral flagella;Costia necatrix(Leclerq) is also 3-flagellate; causes destructive epidemics in fish-hatcheries.

I. RHIZOFLAGELLATA (PANTOSTOMATA)

Food taken in by pseudopodia at any part of the body.

Order 1.—HOLOMASTIGACEAE. Body homaxial with uniform flagella.Multicilia(Cienkowski);Grassia(Fisch, in frog’s blood and gastric mucus).Order 2.—RHIZOMASTIGACEAE. Flagellum 1, 2 or few, diverging from anterior end.Mastigamoeba(F.E. Schulze).

Order 1.—HOLOMASTIGACEAE. Body homaxial with uniform flagella.Multicilia(Cienkowski);Grassia(Fisch, in frog’s blood and gastric mucus).

Order 2.—RHIZOMASTIGACEAE. Flagellum 1, 2 or few, diverging from anterior end.Mastigamoeba(F.E. Schulze).

II. EUFLAGELLATA

Food taken in at one or more definite mouth-spots, or by a true mouth, or by absorption; or nutrition holophytic.

Order 1.—PROTOMASTIGACEAE. Contractile vacuole simple, one or more, or absent; either holozoic, ingesting food by a mouth-spot (or 2 or more), saprophytic, or parasitic.

Family 1.—Oicomonadidae. Flagellum 1, sometimes with a tail-like posterior prominence passing into a temporary flagellum, but without other cytoplasmic processes.Oicomonas(Kent);Cercomonas(Dujardin) (Fig. 1,32, 33);Codonoeca(James-Clark), with a gelatinous theca.Family 2.—Bicoecidae. Differs fromOicomonadidaein a unilateral proboscidiform process next the flagellum; often thecate and stalked, forming branched colonies, like Choanoflagellates in habit.Bicoeca(J.-Cl.),Poteriodendron.Family 3.—Choanoflagellidae(Choanoflagellata, Kent; Craspedomonadina, Stein). As in previous families, but with flagellum surrounded by an obconical or cylindrical rim of cytoplasm, at the base of which is the ingestive area. The cells of this group have the morphology of the flagellate cells (choanocytes) of sponges. They are often colonial, and in the gelatinous colony ofProterospongia, the more internal cells (Fig. 2,15) pass into a definite “reproductive state.” Many stalked forms are epizoic on Entomostracan Crustacea.

Family 2.—Bicoecidae. Differs fromOicomonadidaein a unilateral proboscidiform process next the flagellum; often thecate and stalked, forming branched colonies, like Choanoflagellates in habit.Bicoeca(J.-Cl.),Poteriodendron.

Family 3.—Choanoflagellidae(Choanoflagellata, Kent; Craspedomonadina, Stein). As in previous families, but with flagellum surrounded by an obconical or cylindrical rim of cytoplasm, at the base of which is the ingestive area. The cells of this group have the morphology of the flagellate cells (choanocytes) of sponges. They are often colonial, and in the gelatinous colony ofProterospongia, the more internal cells (Fig. 2,15) pass into a definite “reproductive state.” Many stalked forms are epizoic on Entomostracan Crustacea.

(a) Naked forms often stalked:Monosiga(Kent), stalked solitary;Codosiga(Kent) (Fig. 2,3), stalked social;Desmarella(Kent), unstalked, andAstrosiga(Kent), stalked, form floating colonies.(b) Forms enclosed in a vase-like shell:Salpingoeca(J.-Cl.); (Fig. 2,1, 6, 7) recalling the habit ofMonosigaandCod siga;Polyoecaforming a branched free swimming colony.(c) Forms surrounded by a gelatinous sheath:Proterospongia(Kent) (Fig. 2,15);Phalansterium(Cienk.) (Fig. 1,12), has a slender cylindrical collar, and a branching tubular stalk.

(a) Naked forms often stalked:Monosiga(Kent), stalked solitary;Codosiga(Kent) (Fig. 2,3), stalked social;Desmarella(Kent), unstalked, andAstrosiga(Kent), stalked, form floating colonies.

(b) Forms enclosed in a vase-like shell:Salpingoeca(J.-Cl.); (Fig. 2,1, 6, 7) recalling the habit ofMonosigaandCod siga;Polyoecaforming a branched free swimming colony.

(c) Forms surrounded by a gelatinous sheath:Proterospongia(Kent) (Fig. 2,15);Phalansterium(Cienk.) (Fig. 1,12), has a slender cylindrical collar, and a branching tubular stalk.

Family 4.—Haemoflagellidae. Forms with a complex nuclear apparatus, and a muscular undulating membrane with which one or two flagella are connected, parasitic in Metazoa (often in the blood).Trypanosoma(Gruby) (Fig. 1,21, 22),Herpetomonas(Kent),Treponema(Vuillemin)(=Spirochaete, auctt., nec. Ehrbg.).Family 5.—Amphimonadidae. Flagella 2 anterior, both directed forward, equal and similar; in stalk sheath, &c., often recalling Choanoflagellata,Amphimonas(Kent),Diplomitus(Kent);Spongomonas(St.), with thick branching gelatinous sheath.Family 6.—Monadidae. Flagella 2 (3), anterior all directed forwards, one long the other (or 2) accessory, short.Monas(St.);Anthophysa(Bory) (Fig. 2,12, 13), with the stalk composed of the accumulation of faeces at the hinder end of the cells of the colony.Family 7.—Bodonidae. Flagella 2 (or 3) 1 anterior, the other (1 or 2) antero-lateral and trailing or becoming fixed at the end to form a temporary anchor.Bodo(Ehrb.) (figs. 1,23-26and 2,10).B. lensis the “hooked” andB. saltansthe “springing monad” of Dallinger and Drysdale;Dallingeria(Kent) with a pair of antero-lateral flagella;Costia necatrix(Leclerq) is also 3-flagellate; causes destructive epidemics in fish-hatcheries.

Family 5.—Amphimonadidae. Flagella 2 anterior, both directed forward, equal and similar; in stalk sheath, &c., often recalling Choanoflagellata,Amphimonas(Kent),Diplomitus(Kent);Spongomonas(St.), with thick branching gelatinous sheath.

Family 6.—Monadidae. Flagella 2 (3), anterior all directed forwards, one long the other (or 2) accessory, short.Monas(St.);Anthophysa(Bory) (Fig. 2,12, 13), with the stalk composed of the accumulation of faeces at the hinder end of the cells of the colony.

Family 7.—Bodonidae. Flagella 2 (or 3) 1 anterior, the other (1 or 2) antero-lateral and trailing or becoming fixed at the end to form a temporary anchor.Bodo(Ehrb.) (figs. 1,23-26and 2,10).B. lensis the “hooked” andB. saltansthe “springing monad” of Dallinger and Drysdale;Dallingeria(Kent) with a pair of antero-lateral flagella;Costia necatrix(Leclerq) is also 3-flagellate; causes destructive epidemics in fish-hatcheries.

1.Salpingoeca fusiformis, S. Kent (Choanoflagellata). The protoplasmic body is drawn together within the goblet-shaped shell, and divided into numerous spores.

2. Escape of the spores of the same as monoflagellate and swarm-spores.

3.Codosiga umbellata, Tatem (Choanoflagellata); adult colony formed by dichotomous growth.

4. A single zooid of the same.a= nucleus.b= contractile vacuole.c= the characteristic “collar” of naked streaming protoplasm.

5.Hexamita inflata, Duj.(Distomatidae); normal adult.

6, 7Salpingoeca urceolata, S Kent (Choanoflagellata)—6, with collar extended; 7, with collar retracted within the stalked cup.

8Polytoma uvella, Mull. sp. (Chlamydomonadidae).

9.Lophomonas blattarum, Stein (Trichonymphidae) from the intestine ofBlatta orientalis.

10.Bodolens, Mull. (Bodonidae), the wavy filament is a tractellum, the straight one is a trailing thread.

11.Tetramitus sulcatus, Stein (Tetramitidae)

12.Anthophysa vegetans, O.F. Müller (Monadidae). A typical, erect, shortly-branching colony stock with four terminal monad-clusters.

13. Monad cluster of the same in optical section, showing the relation of the individual monads or flagellate zooids to the stemd.

14.Tetramitus rostratus, Perty (Tetramitidae).a= nucleus.b= contractile vacuole.

15.Proterospongia Haeckeli, Saville Kent (Choanoflagellata); A social colony of about forty flagellate zooids.a= nucleus.b= contractile vacuole.c= amoebiform cell sunk within the colonial gelatinous test compared by S. Kent to a mesoderm cell of the sponges.d= similar cell reproducing by transverse fission.e= normal cells, with their collars contracted.f= substance of test.g= individual reproducing by multiple fission, producing microzoospores, comparable to the spermatozoa of sponges.

1.Trichonympha agilis, Leidy, from gut of White Ant (Termite).

2.Opalina ranarum, Purkinje parasitic in frog rectum multinucleate adult.

3, 4. Binary fissions of same, 1-nucleat individual at final stage of fission.

5. Same encysted dejected from rectum to be swallowed by tadpole.

6. Young 1-nucleate individual emerged from cyst, destined to grow, proliferating its nuclei to adult form.a= nucleus.b= food (?) particles in Fig. 1.

Family 8.—Tetramitidae. Body pyriform, the pointed end posterior; flagella 4 anterior.Tetramitus(Perty) (T. calycinusof Kent, Fig. 2,11, 14), is the “calycine monad” of Dallinger and Drysdale;Trichomonas, Donné, possesses a longitudinal undulating membrane, and is an innocuous human parasite; it is possibly related to Haemoflagellates on one hand and toTrichonymphidaeon the other.Family 9.—Distomatidae. Mouth-spots two, or one, with a distinct construction; flagella symmetrically arranged; nucleus bilobed or geminate.Hexamitus(Duj.) (Fig. 2,5), saprophytic and parasitic;Trepomonas(Duj.), freshwater;Megastoma(Grassi) (=Lambliaof Blanchard), with constricted mouth-spot and blepharoplast (kineto-nucleus) parasitic in the small intestine of Mammals, including Man.Family 10.—Trichonymphidae. Flagella numerous, sometimes accompanied by one or more undulating membranes; cytoplasm highly differentiated; contractile vacuole absent; all parasitic in insects (all exceptLophomonasin Termites—the so-called White Ants.)Lophomonas(St.) (Fig. 2,9); parasitic in the cockroach;Dinenympha(Leidy),Pyrsonympha(Leidy);Trichenympha(Leidy) (Fig. 3,1).Family 11.—Opalinidae. Flagella short, numerous, ciliform. uniformly distributed over the flat oval body; nuclei small, numerous, uniform.Only genus,Opalina(Purkinje and Valentin) (Fig. 3,2-6), in bladder and cloaca of the frog (usually regarded as an aberrant ciliate, but E.R. Lankester expressed doubts as to its position in the 9th edition of this encyclopaedia).Order 2.—CHRYSOMONADACEAE.Contractile vacuole simple (in fresh-water forms) or absent; plastids yellow or brown always present; reserves fat.Family 1.—Chrysomonadidae. Body naked, often amoeboid in active state, or sometimes with a cup-like theca, a gelatinous investment, a firm cuticle, or silicified shell; reserves fat or leucosin (starch inZooxanthella); eye-spot present.Chromulina(Cienk.) often forms a golden scum on tanks;Chrysamoeba(Klebs);Hydrurus(Agardh), theca of colonyforming branching tubes, simulating a yellow Conferva in mountain torrents;Dinobryon(Ehrb.) (Fig. 1,8, 15);Stylochrysalis(St.);Uroglena(Ehrb.);Syncrypta(Ehrb.), andSynura(Ehrb.) (Fig. 1,5) form floating spherical colonies;Zooxanthella(Brandt), symbiotic as “yellow cells” in RadiolariaForaminifera,Millepora, and many Actinozoa.Family 2.—Coccolithophoridae. Body invested in a spherical test strengthened by calcareous elements, tangential circular plates, “coccoliths,” “discoliths,” “cyatholiths,” or radiating rods “rhabdoliths.” These are often found in Foraminiferal ooze and its fossil condition, chalk; when coherent as in the complete test, they are known as “coccospheres” and “rhabdospheres.”Coccolithophora(Lohmann),Rhabdosphaera(Haeckel).Order 3.—CRYPTOMONADACEAE.Contractile vacuole (in freshwater forms) simple; plastids green, more rarely red, brown or absent; reserves starch; holophytic or saprophytic.Cryptomonas(Ehrb.);Paramoeba(Greeff) has yellow plastids and shows two cycles, in the one amoeboid, finally encysting to produce a brood of flagellulae; in the other flagellate, and multiplying by longitudinal fission (it differs fromMastigamoebain possessing no flagellum in the amoeboid state, though it takes in food amoeba-fashion);Chilomonas(Ehrb.).Order 4.—CHLOROMONADACEAE.Contractile vacuoles 1-3, a complex of variable arrangement; pellicle delicate; plastids discoid chlorophyll-bodies; reserves oil; eye-spot absent even in active state; holophytic or saprophytic, though with an anterior blind tubular depression simulating a pharynx.Coelomonas(St.),Vacuolaria(Cienk.).Order 5.—EUGLENACEAE.Vacuole large, a reservoir for one or more accessory vacuoles, contractile and opening to the surface by a canal (“pharynx”) in which are planted one or two strong flagella; pellicle strong often striated; nucleus large, chromatophores green, complex or absent; reserves paramylum granules of definite shape, and oil; nutrition variable; body stiff or “metabolic,” never amoeboid. Among the true Flagellates these are the largest, few being below 40 μ and several attaining 130 μ in length of cell-body (excluding flagellum). Encysted condition common; the green forms sometimes multiply in this state and simulate unicellular Algae.Family 1.—Euglenidae. Radial (monaxial) forms; nutrition saprophytic or holophytic, mostly one flagellate. (1) Chromatophore large; eye-spot conspicuous.Euglena(Ehrb.) (Fig. 1,13, 17), with flexible cuticle and metabolic movements (this is probably Priestley’s “green matter” through which he obtained oxygen gas)—a very common genus;Colacium(Ehbg.), in its resting state epizoic on Copepoda, which it colours green;Eutreptia(Perty), biflagellate;Ascoglena(St.);Trachelomonas(Ehrb.), with a hard brown cuticle;Phacus(Nitszche), with a firm rigid pellicle, often symmetrically flattened;Cryptoglena(Ehbg.). (2) Chromatophores absent.Astasia(Duj.), body metabolic;Menoidium(Perty), body not metabolic, somewhat inflected and crescentic;Sphenomonas(Stein), with a short accessory trailing flagellum in front peeled;Distigma(Ehbg.) (Fig. 1,27, 28), very metabolic, with two unequal flagella and two dark pigment spots.Family 2.—Peranemidae. Bilaterally symmetrical, often creeping, pharynx highly developed, with a firm rod-like skeleton, sometimes protrusible; nutrition saprophytic and holozoic.Peranema(Ehbg.) andUrceolus(Mereschowsky), uni-flagellate creeping, very metabolic.Petalomonas(St.), uni-flagellate flattened with a deep ventral groove, not metabolic;Heteronema(Duj.) andTropidoscyphus(St.), with a small accessory anterior trailing flagellum;Anisonema(Duj.) andEntosiphon(St.), with the trailing flagellum as long as the tractellum or even much longer.Order 6.—VOLVOCACEAE.Contractile vacuole simple anterior; cell always enclosed in a cellulose wall (sometimes gelatinous) perforated by the two (more rarely four, five) diverging anterior flagella; reserves starch; chlorophyll almost always present, except inPolytoma, sometimes masked by a red pigment; nutrition usually holophytic, rarely saprophytic, never holozoic. Brood-division in active state common, radial.Family 1.—Chlamydomonadidae. Cell-wall firm not gelatinous, rarely forming colonies. Fore-end of the body with two or four (seldom five) flagella. Almost always green in consequence of the presence of a very large single chromatophore. Generally a delicate shell-like envelope of membranous consistence. 1 to 2 simple contractile vacuoles at the base of the flagella. Usually one eye-speck. Division of the protoplasm within the envelope may produce four, eight or more new individuals. This may occur in the swimming or in a resting stage. Also by more continuous fission microgametes of various sizes are formed. Conjugation is frequent.Genera.—Chlorangium(Stein), lacking green chlorophyll;Chlorogonium(Ehr.) (Fig. 1,6, 7);Polytoma(Ehr.) (Fig. 2,8);Chlamydomonas(Ehr.) (Fig. 1,1, 2, 3);Haematococcus(Agardh) (=Chlamydococcus, A. Braun, Stein);Protococcus(Conn, Huxley and Martin);Chlamydomonas(Cienkowski), causes red snow and “bloody rain”;Carteria(Diesing), quadri-flagellate;Spondytomorum(Ehrb.), forming floating colonies;Coccomonas(St.);Phacotus(Perty);Zoochlorella(Brandt), is the name given to undetermined Chlamydomonads found multiplying in the resting state within and in symbiotic relation to other Protozoa, to the freshwater sponge,Ephydatia,Hydra viridis, and to the Turbellarian,Convoluta viridis(in which last species the active form has been recognized as aCarteria).Family 2.—Volvocidae. Cell-wall gelatinous; always associated in colonies; cells, as in Family 1. The number of individuals united to form a colony varies very much, as does the shape of the colony. Reproduction by the continuous division of all or of only certain individuals of the colony, resulting in the production of a daughter colony (from each such individual). In some, probably in all, at certain times copulation of the individuals of distinct sexual colonies takes place, without or with a differentiation of the colonies and of the copulating cells as male and female. The result of the copulation is a resting zygospore (also called zygote or oospermo or fertilized egg), which after a time develops itself into one or more new colonies.Genera.—Gonium(O.F. Müller) (Fig. 1,14);Stephanosphaera(Cohn);Pandorina(Bory de Vine);Eudorina(Ehr.);Volvox(Ehr.) (Fig. 1,18, 20).The sexual reproduction of the colonies of the Volvocaceae is one of the most important phenomena presented by the Protozoa. In some families of Flagellata full-grown individuals become amoeboid, fuse, encyst, and then break up into flagellate spores which develop simply to the parental form (Fig. 1,23to26). In theChlamydomonadidaea single adult individual by division produces small individuals, so-called “microgametes.” These conjugate with one another or with similar microgametes formed by other adults (as in Chlorogonium, Fig. 1,7); or more rarely in certain genera a microgamete conjugates with an ordinary individual megagamete. The result in either case is a “zygote,” a cell formed by fusion of two which divides in the usual way to produce new individuals. The microgamete in this case is the male element and equivalent to a spermatozoon; the megagamete is the female and equivalent to an egg-cell. The zygote is a “fertilized egg,” or oosperm. In some colony-building forms we find that only certain cells produce by division microgametes; and, regarding the colony as a multicellular individual, we may consider these cells as testis-cells and their microgametes as spermatozoa.Cystoflagellata(Rhynchoflagellataof E.R. Lankester) andDinoflagellataare scarcely more than subdivisions of Flagellata; but, following O. Bütschli, we describe them separately; the three groups being united into hisMastigophora.Further Remarks on the Flagellates.—Besides the work of special Protozoologists, such as F. Cienkowski, O. Bütschli, F. v. Stein, F. Schaudinn, W. Saville Kent, &c., the Flagellates have been a favourite study with botanists, especially algologists: we may cite N. Pringsheim, F. Cohn, W.C. Williamson, W. Zopf, P.A. Dangeard, G. Klebs, G. Senn, F. Schütt; the reason for this is obvious. They present a wide range of structure, from the simple amoeboid genera to the highly differentiated cells of Euglenaceae, and the complex colonies ofProterospongiaandVolvox. By some they are regarded as the parent-group of the whole of the Protozoa—a position which may perhaps better be assigned to the Proteomyxa; but they seem undoubtedly ancestral to Dinoflagellates and to Cystoflagellates, as well as to Sporozoa, and presumably to Infusoria. Moreover, the only distinction between theChlamydomonadidaeand the true green Algae or Chlorophyceae is that when the former divide in the resting condition, or are held together by gelatinization of the older cell-walls (Palmellastate), they round off and separate, while the latter divide by a “party wall” so as to give rise either to a cylindrical filament when the partitions are parallel and the axis of growth constant (Confervatype), or to a plate of tissue when the directions alternate in a plane. The same holds good for the Chrysomonadaceae and Cryptomonadaceae, so that these little groups are included in all text-books of botany. Again among Fungi, the zoospores of the Zoosporous Phycomycetes (Chytrydiaceae, Peronosporaceae, Saprolegniaceae) have the characters of theBodonidae. Thus in two directions the Flagellates lead up to undoubted Plants. Probably also the Chlamydomonads have an ancestral relation to the Conjugatae in the widest sense, and the Chrysomonadaceae to the Diatomaceae; both groups of obscure affinity, since even the reproductive bodies have no special organs of locomotion. For these reasons the Volvocaceae, Chloromonadaceae, Chrysomonadaceae and Cryptomonadaceae have been united as Phytoflagellates; and the Euglenaceae might well be added to these. It is easy to understand the relation of the saprophytic and the holophytic Flagellates to true plants. The capacity to absorb nutritive matter in solution (as contrasted with the ingestion of solid matter) renders the encysted condition compatible with active growth, and what in holozoic forms is a true hypnocyst, a state in which all functions are put to sleep, is here only a rest from active locomotion, nutrition being only limited by the supply of nutritive matter from without, and—in thecase of holophytic species—by the illumination: this latter condition naturally limits the possible growth in thickness in holophytes with undifferentiated tissues. The same considerations apply indeed to the larger parasitic organisms among Sporozoa, such as Gregarines and Myxosporidia and Dolichosporidia, which are giants among Protozoa.

Family 8.—Tetramitidae. Body pyriform, the pointed end posterior; flagella 4 anterior.

Tetramitus(Perty) (T. calycinusof Kent, Fig. 2,11, 14), is the “calycine monad” of Dallinger and Drysdale;Trichomonas, Donné, possesses a longitudinal undulating membrane, and is an innocuous human parasite; it is possibly related to Haemoflagellates on one hand and toTrichonymphidaeon the other.

Family 9.—Distomatidae. Mouth-spots two, or one, with a distinct construction; flagella symmetrically arranged; nucleus bilobed or geminate.Hexamitus(Duj.) (Fig. 2,5), saprophytic and parasitic;Trepomonas(Duj.), freshwater;Megastoma(Grassi) (=Lambliaof Blanchard), with constricted mouth-spot and blepharoplast (kineto-nucleus) parasitic in the small intestine of Mammals, including Man.

Family 10.—Trichonymphidae. Flagella numerous, sometimes accompanied by one or more undulating membranes; cytoplasm highly differentiated; contractile vacuole absent; all parasitic in insects (all exceptLophomonasin Termites—the so-called White Ants.)Lophomonas(St.) (Fig. 2,9); parasitic in the cockroach;Dinenympha(Leidy),Pyrsonympha(Leidy);Trichenympha(Leidy) (Fig. 3,1).

Family 11.—Opalinidae. Flagella short, numerous, ciliform. uniformly distributed over the flat oval body; nuclei small, numerous, uniform.Only genus,Opalina(Purkinje and Valentin) (Fig. 3,2-6), in bladder and cloaca of the frog (usually regarded as an aberrant ciliate, but E.R. Lankester expressed doubts as to its position in the 9th edition of this encyclopaedia).

Order 2.—CHRYSOMONADACEAE.Contractile vacuole simple (in fresh-water forms) or absent; plastids yellow or brown always present; reserves fat.

Family 1.—Chrysomonadidae. Body naked, often amoeboid in active state, or sometimes with a cup-like theca, a gelatinous investment, a firm cuticle, or silicified shell; reserves fat or leucosin (starch inZooxanthella); eye-spot present.Chromulina(Cienk.) often forms a golden scum on tanks;Chrysamoeba(Klebs);Hydrurus(Agardh), theca of colonyforming branching tubes, simulating a yellow Conferva in mountain torrents;Dinobryon(Ehrb.) (Fig. 1,8, 15);Stylochrysalis(St.);Uroglena(Ehrb.);Syncrypta(Ehrb.), andSynura(Ehrb.) (Fig. 1,5) form floating spherical colonies;Zooxanthella(Brandt), symbiotic as “yellow cells” in RadiolariaForaminifera,Millepora, and many Actinozoa.Family 2.—Coccolithophoridae. Body invested in a spherical test strengthened by calcareous elements, tangential circular plates, “coccoliths,” “discoliths,” “cyatholiths,” or radiating rods “rhabdoliths.” These are often found in Foraminiferal ooze and its fossil condition, chalk; when coherent as in the complete test, they are known as “coccospheres” and “rhabdospheres.”Coccolithophora(Lohmann),Rhabdosphaera(Haeckel).

Family 2.—Coccolithophoridae. Body invested in a spherical test strengthened by calcareous elements, tangential circular plates, “coccoliths,” “discoliths,” “cyatholiths,” or radiating rods “rhabdoliths.” These are often found in Foraminiferal ooze and its fossil condition, chalk; when coherent as in the complete test, they are known as “coccospheres” and “rhabdospheres.”Coccolithophora(Lohmann),Rhabdosphaera(Haeckel).

Order 3.—CRYPTOMONADACEAE.Contractile vacuole (in freshwater forms) simple; plastids green, more rarely red, brown or absent; reserves starch; holophytic or saprophytic.Cryptomonas(Ehrb.);Paramoeba(Greeff) has yellow plastids and shows two cycles, in the one amoeboid, finally encysting to produce a brood of flagellulae; in the other flagellate, and multiplying by longitudinal fission (it differs fromMastigamoebain possessing no flagellum in the amoeboid state, though it takes in food amoeba-fashion);Chilomonas(Ehrb.).Order 4.—CHLOROMONADACEAE.Contractile vacuoles 1-3, a complex of variable arrangement; pellicle delicate; plastids discoid chlorophyll-bodies; reserves oil; eye-spot absent even in active state; holophytic or saprophytic, though with an anterior blind tubular depression simulating a pharynx.Coelomonas(St.),Vacuolaria(Cienk.).Order 5.—EUGLENACEAE.Vacuole large, a reservoir for one or more accessory vacuoles, contractile and opening to the surface by a canal (“pharynx”) in which are planted one or two strong flagella; pellicle strong often striated; nucleus large, chromatophores green, complex or absent; reserves paramylum granules of definite shape, and oil; nutrition variable; body stiff or “metabolic,” never amoeboid. Among the true Flagellates these are the largest, few being below 40 μ and several attaining 130 μ in length of cell-body (excluding flagellum). Encysted condition common; the green forms sometimes multiply in this state and simulate unicellular Algae.

Order 4.—CHLOROMONADACEAE.Contractile vacuoles 1-3, a complex of variable arrangement; pellicle delicate; plastids discoid chlorophyll-bodies; reserves oil; eye-spot absent even in active state; holophytic or saprophytic, though with an anterior blind tubular depression simulating a pharynx.Coelomonas(St.),Vacuolaria(Cienk.).

Order 5.—EUGLENACEAE.Vacuole large, a reservoir for one or more accessory vacuoles, contractile and opening to the surface by a canal (“pharynx”) in which are planted one or two strong flagella; pellicle strong often striated; nucleus large, chromatophores green, complex or absent; reserves paramylum granules of definite shape, and oil; nutrition variable; body stiff or “metabolic,” never amoeboid. Among the true Flagellates these are the largest, few being below 40 μ and several attaining 130 μ in length of cell-body (excluding flagellum). Encysted condition common; the green forms sometimes multiply in this state and simulate unicellular Algae.

Family 1.—Euglenidae. Radial (monaxial) forms; nutrition saprophytic or holophytic, mostly one flagellate. (1) Chromatophore large; eye-spot conspicuous.Euglena(Ehrb.) (Fig. 1,13, 17), with flexible cuticle and metabolic movements (this is probably Priestley’s “green matter” through which he obtained oxygen gas)—a very common genus;Colacium(Ehbg.), in its resting state epizoic on Copepoda, which it colours green;Eutreptia(Perty), biflagellate;Ascoglena(St.);Trachelomonas(Ehrb.), with a hard brown cuticle;Phacus(Nitszche), with a firm rigid pellicle, often symmetrically flattened;Cryptoglena(Ehbg.). (2) Chromatophores absent.Astasia(Duj.), body metabolic;Menoidium(Perty), body not metabolic, somewhat inflected and crescentic;Sphenomonas(Stein), with a short accessory trailing flagellum in front peeled;Distigma(Ehbg.) (Fig. 1,27, 28), very metabolic, with two unequal flagella and two dark pigment spots.Family 2.—Peranemidae. Bilaterally symmetrical, often creeping, pharynx highly developed, with a firm rod-like skeleton, sometimes protrusible; nutrition saprophytic and holozoic.Peranema(Ehbg.) andUrceolus(Mereschowsky), uni-flagellate creeping, very metabolic.Petalomonas(St.), uni-flagellate flattened with a deep ventral groove, not metabolic;Heteronema(Duj.) andTropidoscyphus(St.), with a small accessory anterior trailing flagellum;Anisonema(Duj.) andEntosiphon(St.), with the trailing flagellum as long as the tractellum or even much longer.

Family 2.—Peranemidae. Bilaterally symmetrical, often creeping, pharynx highly developed, with a firm rod-like skeleton, sometimes protrusible; nutrition saprophytic and holozoic.Peranema(Ehbg.) andUrceolus(Mereschowsky), uni-flagellate creeping, very metabolic.Petalomonas(St.), uni-flagellate flattened with a deep ventral groove, not metabolic;Heteronema(Duj.) andTropidoscyphus(St.), with a small accessory anterior trailing flagellum;Anisonema(Duj.) andEntosiphon(St.), with the trailing flagellum as long as the tractellum or even much longer.

Order 6.—VOLVOCACEAE.Contractile vacuole simple anterior; cell always enclosed in a cellulose wall (sometimes gelatinous) perforated by the two (more rarely four, five) diverging anterior flagella; reserves starch; chlorophyll almost always present, except inPolytoma, sometimes masked by a red pigment; nutrition usually holophytic, rarely saprophytic, never holozoic. Brood-division in active state common, radial.

Family 1.—Chlamydomonadidae. Cell-wall firm not gelatinous, rarely forming colonies. Fore-end of the body with two or four (seldom five) flagella. Almost always green in consequence of the presence of a very large single chromatophore. Generally a delicate shell-like envelope of membranous consistence. 1 to 2 simple contractile vacuoles at the base of the flagella. Usually one eye-speck. Division of the protoplasm within the envelope may produce four, eight or more new individuals. This may occur in the swimming or in a resting stage. Also by more continuous fission microgametes of various sizes are formed. Conjugation is frequent.

Genera.—Chlorangium(Stein), lacking green chlorophyll;Chlorogonium(Ehr.) (Fig. 1,6, 7);Polytoma(Ehr.) (Fig. 2,8);Chlamydomonas(Ehr.) (Fig. 1,1, 2, 3);Haematococcus(Agardh) (=Chlamydococcus, A. Braun, Stein);Protococcus(Conn, Huxley and Martin);Chlamydomonas(Cienkowski), causes red snow and “bloody rain”;Carteria(Diesing), quadri-flagellate;Spondytomorum(Ehrb.), forming floating colonies;Coccomonas(St.);Phacotus(Perty);Zoochlorella(Brandt), is the name given to undetermined Chlamydomonads found multiplying in the resting state within and in symbiotic relation to other Protozoa, to the freshwater sponge,Ephydatia,Hydra viridis, and to the Turbellarian,Convoluta viridis(in which last species the active form has been recognized as aCarteria).

Family 2.—Volvocidae. Cell-wall gelatinous; always associated in colonies; cells, as in Family 1. The number of individuals united to form a colony varies very much, as does the shape of the colony. Reproduction by the continuous division of all or of only certain individuals of the colony, resulting in the production of a daughter colony (from each such individual). In some, probably in all, at certain times copulation of the individuals of distinct sexual colonies takes place, without or with a differentiation of the colonies and of the copulating cells as male and female. The result of the copulation is a resting zygospore (also called zygote or oospermo or fertilized egg), which after a time develops itself into one or more new colonies.

Genera.—Gonium(O.F. Müller) (Fig. 1,14);Stephanosphaera(Cohn);Pandorina(Bory de Vine);Eudorina(Ehr.);Volvox(Ehr.) (Fig. 1,18, 20).

The sexual reproduction of the colonies of the Volvocaceae is one of the most important phenomena presented by the Protozoa. In some families of Flagellata full-grown individuals become amoeboid, fuse, encyst, and then break up into flagellate spores which develop simply to the parental form (Fig. 1,23to26). In theChlamydomonadidaea single adult individual by division produces small individuals, so-called “microgametes.” These conjugate with one another or with similar microgametes formed by other adults (as in Chlorogonium, Fig. 1,7); or more rarely in certain genera a microgamete conjugates with an ordinary individual megagamete. The result in either case is a “zygote,” a cell formed by fusion of two which divides in the usual way to produce new individuals. The microgamete in this case is the male element and equivalent to a spermatozoon; the megagamete is the female and equivalent to an egg-cell. The zygote is a “fertilized egg,” or oosperm. In some colony-building forms we find that only certain cells produce by division microgametes; and, regarding the colony as a multicellular individual, we may consider these cells as testis-cells and their microgametes as spermatozoa.

Cystoflagellata(Rhynchoflagellataof E.R. Lankester) andDinoflagellataare scarcely more than subdivisions of Flagellata; but, following O. Bütschli, we describe them separately; the three groups being united into hisMastigophora.

Further Remarks on the Flagellates.—Besides the work of special Protozoologists, such as F. Cienkowski, O. Bütschli, F. v. Stein, F. Schaudinn, W. Saville Kent, &c., the Flagellates have been a favourite study with botanists, especially algologists: we may cite N. Pringsheim, F. Cohn, W.C. Williamson, W. Zopf, P.A. Dangeard, G. Klebs, G. Senn, F. Schütt; the reason for this is obvious. They present a wide range of structure, from the simple amoeboid genera to the highly differentiated cells of Euglenaceae, and the complex colonies ofProterospongiaandVolvox. By some they are regarded as the parent-group of the whole of the Protozoa—a position which may perhaps better be assigned to the Proteomyxa; but they seem undoubtedly ancestral to Dinoflagellates and to Cystoflagellates, as well as to Sporozoa, and presumably to Infusoria. Moreover, the only distinction between theChlamydomonadidaeand the true green Algae or Chlorophyceae is that when the former divide in the resting condition, or are held together by gelatinization of the older cell-walls (Palmellastate), they round off and separate, while the latter divide by a “party wall” so as to give rise either to a cylindrical filament when the partitions are parallel and the axis of growth constant (Confervatype), or to a plate of tissue when the directions alternate in a plane. The same holds good for the Chrysomonadaceae and Cryptomonadaceae, so that these little groups are included in all text-books of botany. Again among Fungi, the zoospores of the Zoosporous Phycomycetes (Chytrydiaceae, Peronosporaceae, Saprolegniaceae) have the characters of theBodonidae. Thus in two directions the Flagellates lead up to undoubted Plants. Probably also the Chlamydomonads have an ancestral relation to the Conjugatae in the widest sense, and the Chrysomonadaceae to the Diatomaceae; both groups of obscure affinity, since even the reproductive bodies have no special organs of locomotion. For these reasons the Volvocaceae, Chloromonadaceae, Chrysomonadaceae and Cryptomonadaceae have been united as Phytoflagellates; and the Euglenaceae might well be added to these. It is easy to understand the relation of the saprophytic and the holophytic Flagellates to true plants. The capacity to absorb nutritive matter in solution (as contrasted with the ingestion of solid matter) renders the encysted condition compatible with active growth, and what in holozoic forms is a true hypnocyst, a state in which all functions are put to sleep, is here only a rest from active locomotion, nutrition being only limited by the supply of nutritive matter from without, and—in thecase of holophytic species—by the illumination: this latter condition naturally limits the possible growth in thickness in holophytes with undifferentiated tissues. The same considerations apply indeed to the larger parasitic organisms among Sporozoa, such as Gregarines and Myxosporidia and Dolichosporidia, which are giants among Protozoa.

Literature.—W.S. Kent,Manual of the Infusoria, vol. i. Protozoa (1880-1882); O. Bütschli,Die Flagellaten(in Bronn’sThierreich, vol. i. Protozoa, 1885); these two works contain full bibliographies of the antecedent authors. See also J. Goroschankin (on Chlamydomonads) inBull. Soc. Nat.(Moscow, iv. v., 1890-1891); G. Klebs, “Flagellatenstudien” inZeitsch. Wiss. Zool.lv. (1892); Doflein,Protozoen als Krankheitserreger(1900); Senn, “Flagellaten,” in Engler and Prantl’sPflanzenfamilien, 1 Teil, Abt. 1a (1900); R. Francé,Der Organismus der Craspedomonaden(1897); Grassi and Sandias, “Trichonymphidae,” inQuart. J. Micr. Sci.xxxix.-xl. (1897); Bezzenberger, “Opa inidae” inArch. Protist, iii. (1903); Marcus Hartog, “Protozoa,” inCambridge Nat. Hist.vol. i. (1906).

(M. Ha.)

FLAGEOLET,in music, a kind offlute-à-becwith a new fingering, invented in France at the end of the 16th century, and in vogue in England from the end of the 17th to the beginning of the 19th century. The instrument is described and illustrated by Mersenne,1who states that the most famous maker and player in his day was Le Vacher. The flageolet differed from the recorder in that it had four finger-holes in front and two thumb-holes at the back instead of seven finger-holes in front and one thumb-hole at the back. This fingering has survived in the French flageolet still used in the provinces of France in small orchestras and for dance music. The arrangement of the holes was as follows: 1, left thumb-hole at the back near mouthpiece; 2 and 3, finger-holes stopped by the left hand; 4, finger-hole stopped by right hand; 5, thumb-hole at the back; 6, hole near the open end. According to Dr Burney (History of Music) the flageolet was invented by the Sieur Juvigny, who played it in theBallet comique de la Royne, 1581. Dr Edward Browne,2writing to his father from Cologne on the 20th of June 1673, relates, “We have with us here one ... and Mr Hadly upon the flagelet, which instrument he hath so improved as to invent large ones and outgoe in sweetnesse all the basses whatsoever upon any other instrument.” About the same time was published Thomas Greeting’sPleasant Companion; or New Lessons and Instructions for the Flagelet(London, 1675 or 1682), a rare book of which the British Museum does not possess a copy. The instrument retained its popularity until the beginning of the 19th century, when Bainbridge constructed double and triple flageolets.3The three tubes were bored parallel through one piece of wood communicating near the mouthpiece which was common to all three. The lowest notes of the respective tubes were

The word flageolet was undoubtedly derived from the medieval Fr.flajol, the primitive whistle-pipe.

(K. S.)

1Harmonie universelle(Paris, 1636), bk. v. pp. 232-237.2See Sir Thomas Browne’s Works, vol. i. p. 206.3See Capt. C.R. Day,Descriptive Catalogue of Musical Instruments(London, 1891), pp. 18-22 and pl. 4; alsoComplete Instructions for the Double Flageolet(London, 1825); andThe Preceptor, or a Key to the Double Flageolet(London, 1815).

1Harmonie universelle(Paris, 1636), bk. v. pp. 232-237.

2See Sir Thomas Browne’s Works, vol. i. p. 206.

3See Capt. C.R. Day,Descriptive Catalogue of Musical Instruments(London, 1891), pp. 18-22 and pl. 4; alsoComplete Instructions for the Double Flageolet(London, 1825); andThe Preceptor, or a Key to the Double Flageolet(London, 1815).

FLAGSHIP,the vessel in a fleet which carries the flag, the symbol of authority of an admiral.

FLAHAUT DE LA BILLARDERIE, AUGUSTE CHARLES JOSEPH,Comte de(1785-1870), French general and statesman, son of Alexandre Sébastien de Flahaut de la Billarderie, comte de Flahaut, beheaded at Arras in February 1793, and his wife Adélaide Filleul, afterwards Mme de Souza (q.v.), was born in Paris on the 21st of April 1785. Charles de Flahaut was generally recognized to be the offspring of his mother’s liaison with Talleyrand, with whom he was closely connected throughout his life. His mother took him with her into exile in 1792, and they remained abroad until 1798. He entered the army as a volunteer in 1800, and received his commission after the battle of Marengo. He became aide-de-camp to Murat, and was wounded at the battle of Landbach in 1805. At Warsaw he met Anne Poniatowski, Countess Potocka, with whom he rapidly became intimate. After the battle of Friedland he received the Legion of Honour, and returned to Paris in 1807. He served in Spain in 1808, and then in Germany. Meanwhile the Countess Potocka had established herself in Paris, but Charles de Flahaut had by this time entered on his liaison with Hortense de Beauharnais, queen of Holland. The birth of their son was registered in Paris on the 21st of October 1811 as Charles Auguste Louis Joseph Demorny, known later as the due de Morny. Flahaut fought with distinction in the Russian campaign of 1812, and in 1813 became general of brigade, aide-de-camp to the emperor, and, after the battle of Leipzig, general of division. After Napoleon’s abdication in 1814 he submitted to the new government, but was placed on the retired list in September. He was assiduous in his attendance on Queen Hortense until the Hundred Days brought him into active service again. A mission to Vienna to secure the return of Marie Louise resulted in failure. He was present at Waterloo, and afterwards sought to place Napoleon II. on the throne. He was saved from exile by Talleyrand’s influence, but was placed under police surveillance. Presently he elected to retire to Germany, and thence to England, where he married Margaret, daughter of Admiral George Keith Elphinstone, Lord Keith, and after the latter’s death Baroness Keith in her own right. The French ambassador opposed the marriage, and Flahaut resigned his commission. His eldest daughter, Emily Jane, married Henry, 4th marquess of Lansdowne. The Flahauts returned to France in 1827, and in 1830 Louis Philippe gave the count the grade of lieutenant-general and made him a peer of France. He remained intimately associated with Talleyrand’s policy, and was, for a short time in 1831, ambassador at Berlin. He was afterwards attached to the household of the duke of Orleans, and in 1841 was sent as ambassador to Vienna, where he remained until 1848, when he was dismissed and retired from the army. After thecoup d’étatof 1851 he was again actively employed, and from 1860 to 1862 was ambassador at the court of St James’s. He died on the 1st of September 1870. The comte de Flahaut is perhaps better remembered for his exploits in gallantry, and the elegant manners in which he had been carefully trained by his mother, than for his public services, which were not, however, so inconsiderable as they have sometimes been represented to be.

See A. de Haricourt,Madame de Souza et sa famille(1907).

FLAIL(from Lat.flagellum, a whip or scourge, but used in the Vulgate in the sense of “flail”; the word appears in Dutchvlegel, Ger.Flegel, and Fr.fléau), a farm hand-implement formerly used for threshing corn. It consists of a short thick club called a “swingle” or “swipple” attached by a rope or leather thong to a wooden handle in such a manner as to enable it to swing freely. The “flail” was a weapon used for military purposes in the middle ages. It was made in the same way as a threshing-flail but much stronger and furnished with iron spikes. It also took the form of a chain with a spiked iron ball at one end swinging free on a wooden or iron handle. This weapon was known as the “morning star” or “holy water sprinkler.” During the panic over the Popish plot in England from 1678 to 1681, clubs, known as “Protestant flails,” were carried by alarmed Protestants (seeGreen Ribbon Club).

FLAMBARD, RANULF,orRalph(d. 1128), bishop of Durham and chief minister of William Rufus, was the son of a Norman parish priest who belonged to the diocese of Bayeux. Migrating at an early age to England, the young Ranulf entered the chancery of William I. and became conspicuous as a courtier. He was disliked by the barons, who nicknamed him Flambard in reference to his talents as a mischief-maker; but he acquired the reputation of an acute financier and appears to have played an important part in the compilation of the Domesday survey. In that record he is mentioned as a clerk by profession, and as holding land both in Hants and Oxfordshire. Before the death of the old king he became chaplain to Maurice, bishop of London, under whom he had formerly served in the chancery. But early in the next reign Ranulf returned to the royal service. He is usually described as the chaplain of Rufus; he seems in that capacity to have been the head of the chancery and the custodian of the great seal. But he is also called treasurer;and there can be no doubt that his services were chiefly of a fiscal character. His name is regularly connected by the chroniclers with the ingenious methods of extortion from which all classes suffered between 1087 and 1100. He profited largely by the tyranny of Rufus, farming for the king a large proportion of the ecclesiastical preferments which wereillegallykept vacant, and obtaining for himself the wealthy see of Durham (1099). His fortunes suffered an eclipse upon the accession of Henry I., by whom he was imprisoned in deference to the popular outcry. A bishop, however, was an inconvenient prisoner, and Flambard soonsucceededin effecting his escape from the Tower of London. A popular legend represents the bishop as descending from the window of his cell by a rope which friends had conveyed to him in a cask of wine. He took refuge with Robert Curthose in Normandy and became one of the advisers who pressed the duke to dispute the crown of England with his younger brother; Robert rewarded the bishop by entrusting him with the administration of the see of Lisieux. After the victory of Tinchebrai (1106) the bishop was among the first to make his peace with Henry, and was allowed to return to his English see. At Durham he passed the remainder of his life. His private life was lax; he had at least two sons, for whom he purchased benefices before they had entered on their teens; and scandalous tales are told of the entertainments with which he enlivened his seclusion. But he distinguished himself, even among the bishops of that age, as a builder and a pious founder. He all but completed the cathedral which his predecessor, William of St Carilef, had begun; fortified Durham; built Norham Castle; founded the priory of Mottisfout and endowed the college of Christchurch, Hampshire. As a politician he ended his career with his submission to Henry, who found in Roger of Salisbury a financier not less able and infinitely more acceptable to the nation. Ranulf died on the 5th of September 1128.

See Orderic Vitalis,Historia ecclesiastica, vols. iii. and iv. (ed. le Prévost, Paris, 1845); the first continuation of Symeon’sHistoria Ecclesiae Dunelmensis(Rolls ed., 1882); William of Malmesbury in theGesta pontificum(Rolls ed., 1870); and thePeterborough Chronicle(Rolls ed., 1861). Of modern writers E.A. Freeman in hisWilliam Rufus(Oxford, 1882) gives the fullest account. See also T.A. Archer in theEnglish Historical Review, ii. p. 103; W. Stubbs’sConstitutional History of England, vol. i. (Oxford, 1897); J.H. Round’sFeudal England(London, 1895).

(H. W. C. D.)

FLAMBOROUGH HEAD,a promontory on the Yorkshire coast of England, between the Filey and Bridlington bays of the North Sea. It is a lofty chalk headland, and the resistance it offers to the action of the waves may be well judged by contrast with the low coast of Holderness to the south. The cliffs of the Head, however, are pierced with caverns and fringed with rocks of fantastic outline. Remarkable contortion of strata is seen at various points in the chalk. Sea-birds breed abundantly on the cliffs. A lighthouse marks the point, in 54° 7′ N., 0° 5′ W.

FLAMBOYANT STYLE,the term given to the phase of Gothic architecture in France which corresponds in period to the Perpendicular style. The word literally means “flowing” or “flaming,” in consequence of the resemblance to the curved lines of flame in window tracery. The earliest examples of flowing tracery are found in England in the later phases of the Decorated style, where, in consequence of the omission of the enclosing circles of the tracery, the carrying through of the foliations resulted in a curve of contrary flexure of ogee form and hence the term flowing tracery. In the minster and the church of St Mary at Beverley, dating from 1320 and 1330, are the earliest examples in England; in France its first employment dates from about 1460, and it is now generally agreed that the flamboyant style was introduced from English sources. One of the chief characteristics of the flamboyant style in France is that known as “interpenetration,” in which the base mouldings of one shaft are penetrated by those of a second shaft of which the faces are set diagonally. This interpenetration, which was in a sense atour de forceof French masons, was carried to such an extent that in a lofty rood-screen the mouldings penetrating the base-mould would be found to be those of a diagonal buttress situated 20 to 30 ft. above it. It was not limited, however, to internal work; in late 15th and early 16th century ecclesiastical architecture it is found on the façades of some French cathedrals, and often on the outside of chapels added in later times.

FLAME(Lat.flamma; the rootflag-appears inflagrare, to burn, blaze, and Gr.φλέγειν). There is no strict scientific definition of flame, but for the purpose of this article it will be regarded as a name for gas which is temporarily luminous in consequence of chemical action. It is well known that the luminosity of gases can be induced by the electrical discharge, and with rapidly alternating high-tension discharges in air an oxygen-nitrogen flame is produced which is long and flickering, can be blown out, yields nitrogen peroxide, and is in fact indistinguishable from an ordinary flame except by its electrical mode of maintenance. The term “flame” is also applied to solar protuberances, which, according to the common view, consist of gases whose glow is of a purely thermal origin. Even with the restricted definition given above, difficulties present themselves. It is found, for example, with a hydrogen flame that the luminosity diminishes as the purity of the hydrogen is increased and as the air is freed from dust, and J.S. Stas declared that under the most favourable conditions he was only able, even in a dark room, to localize the flame by feeling for it, an observation consistent with the fact that the line spectrum of the flame lies wholly in the ultra-violet. On the other hand, there are many examples of chemical combination between gases where the attendant radiation is below the pitch of visibility, as in the case of ethylene and chlorine. It will be obvious from these facts that a strict definition of flame is hardly possible. The common distinction between luminous and non-luminous flames is, of course, quite arbitrary, and only corresponds to a rough estimate of the degree of luminosity.

The chemical energy necessary for the production of flame may be liberated during combination or decomposition. A single substance like gun-cotton, which is highly endothermic and gives gaseous products, will produce a bright flame of decomposition if a single piece be heated in an evacuated flask. Combination is the more common case, and this means that we have two separate substances involved. If they be not mixeden massebefore combination, the one which flows as a current into the other is called conventionally the “combustible,” but the simple experiment of burning air in coal gas suffices to show the unreality of this distinction between combustible and supporter of combustion, which, in fact, is only one of the many partial views that are explained and perhaps justified by the dominance of oxygen in terrestrial chemistry.

Although hydrocarbon flames are the commonest and most interesting, it will be well to consider simpler flames first in order to discuss some fundamental problems. In hydrocarbon flames the complexity of the combustible, its susceptibility to change by heating, and the possibilities of fractional oxidation, create special difficulties. In the flame of hydrogen and oxygen or carbon monoxide and oxygen we have simpler conditions, though here, too, things may be by no means so simple as they seem from the equations 2H2+ O2= 2H2O and 2CO + O2= 2CO2. The influence of water vapour on both these actions is well known, and the molecular transactions may in reality be complicated. We shall, however, assume for the sake of clearness that in these cases we have a simple reaction taking place throughout the mass of flame. There are various ways in which a pair of gases may be burned, and these we shall consider separately. Let us first suppose the two gases to have been mixeden masseand a light to be applied to the stationary mixture. If the mixture be made within certain limiting proportions, which vary for each case, a flame spreads from the point where the light is applied, and the flame traverses the mixture. This flame may be very slow in its progress or it may attain a velocity of the order of one or two thousand metres per second. Until comparatively recent times great misunderstanding prevailed on this subject. The slow rate of movement of flame in short lengths of gaseous mixtures was taken to be the velocity of explosion, but more recent researches by M.P.E. Berthelot,E. Mallard and H.L. le Chatelier and H.B. Dixon have shown that a distinction must be made between the slowinitial rate of inflammationof gaseous mixtures and therapid rate of detonation, or rate of theexplosive wave, which in many cases is subsequently set up. We shall here deal only with the slow movements of flame. The development of a flame in such a gaseous mixture requires that a small portion of it should be raised to a temperature called thetemperature of ignition. Here again considerable misunderstanding has prevailed. The temperature of ignition has often been regarded as the temperature at which chemical combination begins, whereas it is really the temperature at which combination has reached a certain rate. The combination of hydrogen and oxygen begins at temperatures far below that of ignition. It may indeed be supposed that the combination occurs with extreme slowness even at ordinary temperatures, and that as the temperature is raised the velocity of the reaction increases in accordance with the general expression according to which an increase of 10°C. will approximately double the rate. However that may be, it has been proved experimentally by J.H. van’t Hoff, Victor Meyer and others that the combination of hydrogen and oxygen proceeds at perceptible rates far below the temperature of ignition. The phenomenon appears to be greatly influenced by the solid surfaces which are present; thus in a plain glass vessel the combination only began to be perceptible at 448°, whilst in a silvered glass vessel it would be detected at 182°C.

The same kind of thing is true for most oxidizable substances, including ordinary combustibles. We must look upon the application of heat to a combustible mixture as resulting in an increase of the rate of combination locally. Let us suppose that we are dealing with a stratum of the mixture in small contiguous sections. If we raise the temperature of the first sectiona°C., an increased rate of combination is set up. The heat produced by this combination will be dissipated by conduction and radiation, and we will suppose that it does not quite suffice to raise the adjacent section of the mixture toa°C. The combination in that section, therefore, will not be as rapid as in the first one, and so evidently the impulse to combination will go on abating as we pass along the stratum. Suppose now we start again and heat the first section of the mixture to a temperaturec°C., such that the rate of combination is very rapid and the heat developed by combination suffices to raise the adjacent section of the mixture to a temperature higher thanc°C. The rate of combination will then be greater than in the first section, and the impulse to combination will be intensified in the same way from section to section along the stratum until a maximum temperature is reached. It is obvious that there must be a temperature ofb°C. betweena° andc° which will satisfy this condition, that the heat which results from the combination stimulated in the first section just suffices to raise the temperature of the second section tob°. This temperatureb° is the temperature of ignition of the mixture; so soon as it is attained by a portion of the mixture the combustion becomes self-sustaining and flame spreads through the mixture. Ignition temperature may be defined briefly as the temperature at which the initial loss of heat due to conduction, &c., is equal to the heat evolved in the same time by the chemical reaction (van’t Hoff). From the above considerations we see that the temperature of ignition will vary not only when the gases are varied, but when the proportions of the same gases are varied, and also when the pressure is varied. We can see also that outside certain limiting proportions a mixture of gases will have no practicable ignition temperature, that is to say, the cooling effect of the gas which is in excess will carry off so much heat that no attainable initial heating will suffice to set up the transmission of a constant temperature. Thus in the case of hydrogen and air, mixtures containing less than 5 and more than 72% of hydrogen are not inflammable. The theory of ignition temperature enables us to understand why in an explosive mixture a very small electric spark may not suffice to induce explosion. Combination will indeed take place in the path of the spark, but the amount of it is not sufficient to meet the loss of heat by conduction, &c. It must be added that the theory of ignition temperatures given above does not explain all the observed facts. F. Emich states that the inflammability of gaseous mixtures is not necessarily greatest when the gases are mixed in the proportions theoretically required for complete combination, and the influence of foreign gases does not appear to follow any simple law. The presence of a small quantity of a gas may exercise a profound influence on the ignition temperature as in the case of the addition of ethylene to hydrogen (Sir Edward Frankland), and again when a mixture of methane and air is raised to its ignition temperature a sensible interval (about 10 seconds) elapses before inflammation occurs.

The rate at which a flame will traverse a mixture of two gases which has been ignited depends on the proportions in which the gases are mixed. Fig. 1 (Bunte) represents this relationship for several common gases.

If a ready-made gaseous mixture is to be used for the production of a steady flame, it may be forced through a tube and ignited at the end; it is obvious that the velocity of efflux must be greater than the initial rate of inflammation of the mixture, for otherwise the mixture would fire back down the tube. If the velocity of efflux be considerably greater than the rate of inflammation, the flame will be separated from the end of the tube, and only appear as a flickering crown where the velocity and inflammability of the issuing gas have been diminished by admixture with air. With much increased velocity of efflux the flame will be blown out. J.B.A. Dumas used to show the experiment of blowing out a candle with electrolytic gas. A steady flame formed by burning a ready-made gaseous mixture at the end of a tube of circular section has the form shown in fig. 2. The small internal cone marks the lower limiting surface of the flame; it is the locus of all points where the velocity of efflux is just equal to the velocity of inflammation, and its conical form is explained by the fact that the rate of efflux of gas is greatest in the vertical axis of the tube where the flow is not retarded by friction with the walls, as well as by the further fact that the gas issuing from such an orifice spreads outwards, the inflammation proceeding directly against it. The flame, it will be seen, is of considerable thickness. If the gaseous mixture be hydrogen and oxygen, or carbon monoxide and oxygen, it will have no obvious features of structure beyond those shown in the figure; that is to say, the shaded region of burning gas has the appearance of homogeneity and uniform colour which might be expected to accompany a uniform chemical condition. Some admixture of the external air will, of course, take place, especially in the upper parts of the flame, and detectable quantities of oxides of nitrogen may be found in the products of combustion, but this is an inconsiderable feature. The flame just described is essentially that of a blowpipe.

A second way of producing a flame is the more common one of allowing one gas to stream into the other. Using the same gases as before, hydrogen or carbon monoxide with oxygen, we findagain that the flame is conical in form and uniform in colour, but in this case, if the velocity of efflux be not immoderate, the burning gas only extends over a comparatively thin shell, limited on the inside by the pure combustible and on the outside by a mixture of the products of combustion with oxygen. The combustible gas has to make its own inflammable mixture with the circumambient oxygen, and we may suppose the column of gas to be burned through as it ascends. The core of unburned gas thus becomes thinner as it ascends and the flame tapers to a point. The external surface of a flame of this kind will for the same consumption of gas be larger than that of a flame where the ready-made mixture of gases is used. If a jet of one gas be sent with a sufficient velocity into another, turbulent admixture takes place and an unsteady sheet of flame of uniform colour is obtained.

A third way of forming a flame is to allow the whole of one gas, mixed with a less quantity of the second than is sufficient for complete combustion, to issue into an atmosphere of the second. This is the case with what are generally known as atmospheric burners, of which the Bunsen burner is the prototype. The development of a flame of this kind can be well studied in the case of carbon monoxide and air. The carbon monoxide is fed into a Bunsen burner with closed air-valve, the burner-tube being prolonged by affixing a glass tube to it by means of a cork. The flame consists of a single conical blue sheet. If now the air-valve be opened very slightly, an internal cone of the same blue colour makes its appearance. The air which has entered through the air-valve (“primary” air) has become mixed with the carbon monoxide and so oxidizes its quota in an internal cone, the rest of the carbon monoxide (diluted now, of course, with carbon dioxide and nitrogen) wandering into the external atmosphere to burn (with “secondary” air) in a second cone. The existence of the internal cone and the subsequent thermal effect lead to slight convexity of surface in the outer cone. If the quantity of primary air be increased more internal combustion can take place. This, however, does not lead to an enlargement of the inner cone, for the increase of air increases the rate of inflammation of the mixture, and the inner cone (which only maintains its stability because the rate of efflux of the mixture is greater than the velocity of inflammation) contracts, and will, as the proportion of primary air is increased, soon evince a tendency to enter the burner-tube. At this stage an interesting phenomenon is to be noticed. When we have reached the point of aeration where the velocity of inflammation of the mixture just surpasses the velocity of efflux, the inner cone enters the burner-tube as a disk and descends, but this downward motion checks the suction flow of air through the valve at the base of the burner, whilst it does not appreciably check the pressure flow of the carbon monoxide through the gas nozzle. The result is that a stratum of gas-mixture poor in air, and therefore of low rate of inflammation, is formed, and when the descending disk of flame meets it, the descent is arrested and the disk returns to the top of the tube, reproducing the inner cone. The full air suction is now restored and the course of events is repeated. This oscillatory action can be maintained almost indefinitely long if the pressure and other conditions be maintained constant. With still more primary air the inner cone of flame simply fires back to the burner nozzle, or, in the last stage, we may have enough air entering to produce a flame of the blast blowpipe type, namely, one where the carbon monoxide mixed with anexcessof primary air burns with a single cone in a steady flame.

By means of a simple contrivance devised by A. Smithells a two-coned flame of the kind described may be resolved into its components. The apparatus is like a half-extended telescope made of two glass tubes, and it is evident that the velocity of a mixture of gases flowing through it must be greater in the narrow tube than in the wider one. If the end of the narrower tube be fixed to a Bunsen burner and the flame be formed at the end of the wider one, then when the air-supply is increased to a certain point the inner cone will descend into the wide tube and attach itself to the upper end of the narrower one. This occurs when the velocity of inflammation is just greater than the upward velocity of the gaseous stream in the wide tube and less than the upward velocity in the narrow tube. If the outer tube be now drawn down, a two-coned flame burns at the end of the inner tube; if the outer tubebeslid up again, it detaches the outer cone and carries it upward. This apparatus has been of use in investigating the progress of combustion in various flames.

Temperature of Flames.—The term “flame-temperature” is used very vaguely and has no clear meaning unless qualified by some description. Itisleast ambiguous when used in reference to flames where the combining gases are mixed in theoretical proportions before issuing from the burner. The flame in such a case has considerable thickness and uniformity, and, though the temperature is not constant throughout, flames of this type given by different combustibles admit of comparison. In other flames where the shells of combustion are thin and envelop large regions of unburned or partly-burned gas, it is not clear how temperature should be specified. An ordinary gas-flame will not, from the point of view of the practical arts, give a sufficient temperature for melting platinum, yet a very thin platinum wire may be melted at the edge of the lower part of such a flame. The maximum temperature of the flame is therefore not in any serious sense an available temperature. It will suffice to point out here that in order to burn a gas so that it may have the highest available temperature, we must burn it with the smallest external flame-surface obtainable. This is done when the combining gases are completely mixed before issuing from the burner. Where this is impracticable we may employ a burner of the Bunsen type, and arrange matters so that a large amount of primary air is supplied. It is in this direction that modern improvements have been made with a view to obtaining hot flames for heating the Welsbach mantle. The Kern burner, for example, employs the principle of the Venturi tube. Where much primary air is drawn in it is usual to provide for it being well mixed with the gas, otherwise an unsteady flame may be produced with a great tendency to light back. The burner head is therefore usually provided with a mixing chamber and the mixture issues through a slit or a mesh. A great many modified Bunsen burners have been produced, the aim in all of them being to produce a flame which shall combine steadiness with the smallest attainable external surface.

To estimate the temperature of flames several methods have been employed. The method of calculation, based on the supposition that the whole heat of combustion is localized in the product (or products) of combustion and heats it to a temperature depending on its specific heat, cannot be applied in a simple way. Apart from the assumption (which there is reason to suppose incorrect) that none of the chemical energy assumes the radiant form directly, we have to regard the possible change of specific heat at high temperatures, the likelihood of dissociation and the time of reaction. Any practical consideration of temperature must have regard to a large assemblage of molecules and not to a single one, and therefore any influence which means delay in combination will result in reduction of temperature by radiation and conduction. It can hardly be maintained that in the present state of knowledge we have the requisite data for the calculation of flame temperature, though good approximations may be made. Many attempts have been made to determine flame temperatures by means of thermo-electric couples and by radiation pyrometers. The couple most employed is that known as H.L. le Chatelier’s, consisting of two wires, one of platinum and the other an alloy of 90% platinum and 10% of rhodium. When all possible precautions are taken it is possible by means of such thermo-couples to measure local flame temperatures with a considerable degree of accuracy. Subjoined are some results obtained at different times and by different observers with regard to the maximum temperatures of flames:—

The following are given by Féry:—

Source of Light in Flames.—We may consider first those flames where solid particles are out of the question; for example, the flame of carbon monoxide in air. The old idea that the luminosity was due to the thermal glow of the highly heated product of combustion has been challenged independently by a number of observers, and the view has been advanced that the emission of light is due to radiation attendant upon a kind of discharge of chemical energy between the reacting molecules. E. Wiedemann proposed the name “chemi-luminescence” for radiation of this kind. The fact is that colourless gases cannot be made to glow by any purely thermal heating at present available, and products of combustion heated to the average temperature of the flames in which they are produced are non-luminous. On the other hand, it must be remembered that in a mass of burning gas only a certain proportion of the molecules are engaged at one instant in the act of chemical combination, and that the energy liberated in such individual transactions, if localized momentarily as heat, would give individual molecules a unique condition of temperature far transcending that of the average, and the distribution of heat in a flame would be very different from that existing in the same mixture of gases heated from an external source to the same average temperature. The view advocated by Smithells is that in the chemical combination of gases the initial phase of the formation of the new molecule is a vibratory one, which directly furnishes light, and that the damping down of this vibration by colliding molecules is the source of that translatory motion which is evinced as heat. This, it will be seen, is an exact reversal of the older view.

The view of Sir H. Davy that “whenever a flame is remarkably brilliant and dense it may always be concluded that some solid matter is produced in it” can be no longer entertained. The flames of phosphorus in oxygen and of carbon disulphide in nitric oxide contain only gaseous products, and Frankland showed that the flames of hydrogen and carbon monoxide became highly luminous under pressure. From his experiments Frankland was led to the generalization that high luminosity of flames is associated with high density of the gases, and he does not draw a distinction in this respect between high density due to high molecular weight and high density due to the close packing of lighter molecules. The increased luminosity of a compressed flame is not difficult to understand from the kinetic theory of gases, but no explanation has appeared of the luminosity considered by Frankland to be due merely to high molecular weight. It is possible that the electron theory may ultimately afford a better understanding of these phenomena.

Structure of Flame.—The vagueness of the term structure, as applied to flames, is to be seen from the very conflicting accounts which are current as to the number of differentiated parts in different flames. Unless this term is restricted to sharp differences in appearance, there is no limit to the number of parts which may be selected for mention. The flame of carbon monoxide, when the gas is not mixed with air before it issues from the burner, shows no clearly differentiated structure, but is a shell of blue luminosity of shaded intensity—a hollow cone if the orifice of the burner be circular and the velocity of the gas not immoderate, or a double sheet of fan shape if the burner have a slit or two inclined pores which cause the jets of issuing gas to spread each other out. Such a flame has but one single distinct feature, and this is not surprising, as there is no reason to suppose that there is any difference in the chemical process or processes that are occurring in different quarters of the flame. The amount of materials undergoing this transformation in different parts of the flame may and does vary; the gases become diluted with products of combustion, and the molecular vibrations gradually die down. These things may cause a variation in the intensity of the light in different quarters, but the differences induced are not sharp or in any proper sense structural. A flame of this kind may develop a secondary feature of structure. If carbon monoxide be burnt in oxygen which is mixed or combined with another element there may be an additional chemical process that will give light; flames in air are sometimes surrounded by a faintly luminous fringe of a greenish cast, apparently associated with the combination of nitrogen with oxygen (H.B. Dixon). Carbon monoxide on being strongly heated begins to dissociate into carbon and carbon dioxide; if the unburnt carbon monoxide within a flame of that gas were so highly heated by its own burning walls as to reach the temperature of dissociation, we might expect to see a special feature of structure due to the separated carbon. Such a temperature does not, however, appear to be reached.

Apart from hydrocarbon flames not much has been published in reference to the structure of flames. The case of cyanogen is of peculiar interest. The beautiful flame of this gas consists of an almost crimson shell surrounded by a margin of bright blue. Investigations have shown that these two colours correspond to two steps in the progress of the combustion, in the first of which the carbon of the cyanogen is oxidized to carbon monoxide and in the second the carbon monoxide oxidized to carbon dioxide.

The inversion of combustion may bring new features of structure into existence; thus when a jet of cyanogen is burnt in oxygen no solid carbon can be found in the flame, but when a jet of oxygen is burnt in cyanogen solid carbon separates on the edge of the flame.

Hydrocarbon Flames.—As already stated the flames of carbon compounds and especially of hydrocarbons have been much more studied than any other kind, as is natural from their common use and practical importance. The earliest investigations were made with coal gas, vegetable oils and tallow, and the composite and complex nature of these substances led to difficulties and confusion in the interpretation of results. One such difficulty may be illustrated by the fact, often overlooked, that when a mixed gaseous combustible issues into air the individual component gases will separate spontaneously in accordance with their diffusibilities: hydrogen will thus tend to get to the outer edge of a flame and heavy hydrocarbons to lag behind.

The features of structure in a hydrocarbon flame depend of course on the manner in which the air is supplied. The extreme cases are (i.) when the issuing gas is supplied before it leaves the burner with sufficient air for complete combustion, as in the blast blowpipe, in which case we have a sheet of blue undifferentiated flame; and (ii.) when the gas has to find all the air it requires after leaving the burner. The intermediate stage is when the issuing gas is supplied before leaving the burner with a part of the air that is required. In this case a two-coned flame is produced. The general theory of such phenomena has already been discussed. It must be remarked that the transition of one kind of flame into the others can be effected gradually, and this is seen with particular ease and distinctness by burning benzene vapour admixed with gradually increasing quantities of air. The key to the explanation of the structure of an ordinary luminous flame, such as that of a candle, is to be found, according to Smithells, by observing the changes undergone by a well-aerated Bunsen flame as the “primary” air is gradually cut off by closing the air-ports at the base of the burner. It is then seen that the two cones of flame evolve or degenerate into the two recognizable blue parts of an ordinary luminous flame, whilst the appearance of the bright yellow luminous patch becomes increasingly emphasized as a hollow dome lying within the upper part of the blue sheath. There are thus three recognizable features of structure in an ordinary luminous flame, each region being as it were a mere shell and the interior of the flame filled with gas which has not yet entered into active combustion. If, as is suggested, the blue parts of an ordinary luminous flame are the relics of the two cones of a Bunsen flame, the chemistry of a Bunsen flame may be appropriately considered first. What happens chemically when a hydrocarbon is burned in a Bunsen burner? The air sent in with the gas is insufficient for completecombustion so that the inner cone of the flame may be considered as air burning in an excess of coal gas. What will be the products of this combustion? This question has been answered at different times in very different ways. There are many conceivable answers: part of the hydrocarbon might be wholly oxidized and the rest left unaltered to mix with the outside air and burn as the outer cone; on the other hand, there might be (as has been so commonly assumed) a selective oxidation in the inner cone whereby the hydrogen was fully oxidized and the carbon set free or oxidized to carbon monoxide; or again the carbon might be oxidized to carbon dioxide or monoxide and the hydrogen set free. There might of course be other intermediate kinds of action. Now it is important at this point to insist upon a distinction between what can be found by direct analysis as to the products of partial combustion, and what can be imagined or inferred as the transitory existence of substances of which the products actually found in analysis are the outcome. We shall consider only in the first instance what substances are found by analysis. Earlier experiments on the Bunsen burner in which coal gas was used, and the gases withdrawn directly from the flame by aspiration, gave no very clear results, but the introduction of the cone-separating apparatus and the use of single hydrocarbons led to more definite conclusions. The analysis of the inter-conal gases from an ethylene flame gave the following numbers:—carbon dioxide = 3.6; water = 9.5; carbon monoxide = 15.6; hydrocarbons = 1.3; hydrogen = 9.4; nitrogen = 60.6.

It appears therefore, and it may be stated as a fact, that a considerable amount of hydrogen is left unoxidized, whilst practically all the carbon is converted into monoxide or dioxide. As the gases have cooled down before analysis and as the reaction CO + H2O ⇄ CO2+ H2is reversible, it may be objected that the inter-conal gases may have a composition when they are hot very different from what they show when cold. Experiments made to test this question have not sustained the objection. Subsequent experiments on the oxidation of hydrocarbons have made it appear undesirable to use the expression “preferential combustion” or “selective combustion” in connexion with the facts just stated; but for the purpose of describing in brief the chemistry of a hydrocarbon flame it is necessary to say that in the inner cone of a Bunsen flame hydrogen and carbon monoxide are the result of the limited oxidation, and that the combustion of these gases with the external air generates the outer cone of the flame. As to the actual stages in the limited oxidation of a hydrocarbon a large amount of very valuable work has been carried out by W.A. Bone and his collaborators. Different hydrocarbons mixed with oxygen have been circulated continuously through a vessel heated to various temperatures, beginning with that (about 250° C.) at which the rate of oxidation is easily appreciable. Proceeding in this way, Bone, without effecting a complete transformation of the hydrocarbon into partially oxidized substances, has isolated large quantities of such products, and concludes that the oxidation of a hydrocarbon involves nothing in the nature of a selective or preferential oxidation of either the hydrogen or the carbon. He maintains that it occurs in several well-defined stages during which oxygen enters into and is incorporated with the hydrocarbon molecule, forming oxygenated intermediate products among which are alcohols and aldehydes. The reactions between ethane and ethylene with an equal volume of oxygen would be represented as follows:—

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