The Works of Francis Balfour 3-14

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I have decided to take early retirement in September 2020. During the many years online I have received wonderful feedback from many readers, researchers and students interested in human embryology. I especially thank my research collaborators and contributors to the site. The good news is Embryology will remain online and I will continue my association with UNSW Australia. I look forward to updating and including the many exciting new discoveries in Embryology!

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Cephalochorda | Urochorda | Elasmobranchii | Teleostei | Cyclostomata | Ganoidei | Amphibia | Aves | Reptilia | Mammalia | Comparison of the Formation of Germinal Layers and Early Stages in Vertebrate Development | Ancestral form of the Chordata | General Conclusions | Epidermis and Derivatives | The Nervous System | Organs of Vision | Auditory, Olfactory, and Lateral Line Sense Organs | Notochord, Vertebral Column, Ribs, and Sternum | The Skull | Pectoral and Pelvic Girdles and Limb Skeleton | Body Cavity, Vascular System and Glands | The Muscular System | Excretory Organs | Generative Organs and Genital Ducts | The Alimentary Canal and Appendages in Chordata
Online Editor 
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This historic 1885 book edited by Foster and Sedgwick is the third of Francis Balfour's collected works published in four editions. Francis (Frank) Maitland Balfour, known as F. M. Balfour, (November 10, 1851 - July 19, 1882) was a British biologist who co-authored embryology textbooks.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
Modern Notes:

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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Draft Version - Notice removed when completed.

Vol. III. A Treatise on Comparative Embryology 2 (1885)


IN many of the Ccelenterata the outermost layer of the blastoderm is converted as a whole into the skin or ectoderm. The cells composing it become no doubt in part differentiated into muscular elements and in part into nervous elements, &c. ; but still it may remain through life as a simple external membrane. This membrane contains in itself indefinite potentialities for developing into various organs, and in all the true Triploblastica these potentialities are more or less realized. The embryonic epiblast ceases in fact, in the higher forms, to become converted as a whole into the epidermis, but first gives rise to parts of the nervous system, organs of special sense, and other parts.

After the formation of these parts the remnant of the epiblast gives rise to the epidermis, and often unites more or less intimately with a subjacent layer of mesoblast, known as the dermis, to form with it the skin.

Various differentiations may arise in the epidermis forming protective or skeletal structures, terminal sense organs, or glands. The structure of the epidermis itself varies greatly, and for Vertebrates its general modifications have been already sufficiently dealt with in chapter XII. Of its special differentiations those of a protective or skeletal nature and those of a glandular nature may be considered in this place.

Protective epidermal structures. These structures constitute a general cuticle or an exoskeleton of scales, hairs, feathers, nails, hoofs, &c. They may be entirely formed from


the epidermis either as (i) a cuticular deposit, or as (2) a chitinization, a cornification, or calcification of its constituent cells. These two processes run into each other, and are in many cases not easily distinguished. The protective structures of the epidermis may be divided into two groups according as they are formed on the outer or the inner side of the epidermis. Dermal skeletal structures are in many cases added to them. Amongst the Invertebrata the most widely distributed type of exoskeleton is a cuticle formed on the outer surface of the epidermis, which reaches its highest development in the Arthropoda. In the same class with this cuticle must be placed the molluscan and brachiopod shells, which are developed as cuticular plates on special regions of the epidermis. They differ, however, from the more usual form of cuticle in their slighter adhesion to the subjacent epidermis, and in their more complicated structure. The test of Ascidians is an abnormal form of exoskeleton belonging to this type. It is originally formed (Hertvvig and Semper) as a cuticle on the surface of the epidermis ; but subsequently epidermic cells migrate into it, and it then constitutes a tissue similar to connective tissue, but differing from ordinary epidermic cuticles in that the cells which deposit it do so over their whole surface, instead of one surface, as is usually the case with epithelial cells.

In the Vertebrata the two types of exoskeleton mentioned above are both found, but that developed on the inner surface of the epidermis is always associated with a dermal skeleton, and that on the outer side frequently so. The type of exoskeleton developed on the inner side of the general epidermis is confined to the Pisces, where it appears as the scales; but a primitive form of these structures persists as the teeth in the Amphibia and Amniota. The type developed on the outer side of the epidermis is almost entirely 1 confined to the Amphibia and Amniota, where it appears as scales, feathers, hairs, claws, nails, &c. For the histological details as to the formation of these various organs I must refer the reader to treatises on histology, confining my attention here to the general embryological processes which take place in their development.

1 The horny teeth of the Cyclostomala are structures belonging to this group.



The most primitive form of the first type of dermal structures is that of the placoid scales of Elasmobranchii 1 . These consist, when fully formed, of a plate bearing a spinous projection. They are constituted of an outer enamel layer on the projecting part, developed as a cuticular deposit of the epidermis (epiblast), and an underlying basis of dentine (the lower part of which may be osseous) with a vascular pulp in its axis. The development (fig. 235) is as follows (Hertwig, No. 306). A papilla of the dermis makes its appearance, the outer layer of which gradually calcifies to form the dentine and osseous tissue. This papilla is covered by the columnar mucous layer of the epidermis (e), from which it is separated by a basement membrane, itself a product of the epidermis. This membrane gradually thickens and calcifies, and so gives rise to the enamel cap (o). The spinous point gradually forces its way through the epidermis, so as to project freely at the surface.

The scales of other forms of fishes are to be derived from those of Elasmobranchii. The great dermal plates of many fishes have been formed by the concrescence of groups of such scales. The dentine in many cases partially or completely atrophies, leaving the major part of the scale formed of osseous tissue ; such plates often become parts of the internal skeleton.




E. epidermis ; C. layers of dermis ; d. uppermost layer of dermis ; p. papilla of dermis ; e. mucous layer of epidermis ; o. enamel layer.

1 For the most important contributions on this subject from which the facts and views here expressed are largely derived, vide O. Hertwig, Nos. 306 808.


The teeth, as will be more particularly described in the section on the alimentary tract, are formed by a modification of the same process as the placoid scales, in which a ridge of the epithelium grows inwards to meet a connective tissue papilla, so that the development of the teeth takes place entirely below the superficial layer of epidermis.

In most Teleostei the enamel and dentine layers have disappeared, and the scales are entirely formed of a peculiar calcified tissue developed in the dermis.

The cuticle covering the scales of Reptiles is the simplest type of protective structure formed on the outer surface of the epidermis. The scales consist of papillae of the dermis and epidermis ; and are covered by a thickened portion of a twolayered cuticle, formed over the whole surface of the body from a cornification of the superficial part of the epidermis. Dermal osseous plates may be formed in connection with these scales, but are never of course united with the superficial cuticle.

Feathers are probably special modifications of such scales. They arise rom an induration of the epidermis of papillae containing a vascular core. The provisional down, usually present at the time of hatching, is formed by the cornification of longitudinal ridges of the mucous layer of the epidermis of the papillee ; each cornified ridge giving rise to a barb of the feather. The horny layer of the epidermis forms a provisional sheath for the developing feather below. When the barbs are fully formed this sheath is thrown- off, the vascular core dries up, and the barbs become free except at their base.

Without entering into the somewhat complicated details of the formation of the permanent feathers, it may be mentioned that the calamus or quill is formed by a cornification in the form of a tube of both layers of the epidermis at the base of the papilla. The quill is open at both ends, and to it is attached the vexillum or plume of the feather. In a typical feather this is formed at the apex of the papilla from ridge-like thickenings of the mucous layer of the epidermis, arranged in the form of a longitudinal axis, continuous with the cornified mucous layer of the quill, and from lateral ridges. These subsequently become converted into the axis and barbs of the plume. The external epidermic layer becomes converted into a provisional horny sheath for the true feather beneath.

On the completion of the plume of the feather the external sheath is thrown off, leaving it quite free, and the vascular core belonging to it shrivels up. The papilla in which the feather is formed becomes at a very early period secondarily enveloped in a pit or follicle which gradually deepens as the development of the feather is continued.

Hairs (Kolliker, No. 298) are formed in solid processes of the mucous layer of the epidermis, which project into the


subjacent dermis. The hair itself arises from a cornification of the cells of the axis of one of the above processes ; and is invested by a sheath similarly formed from the more superficial epidermic cells. A small papilla of the dermis grows into the inner end of the epidermic process when the hair is first formed. The first trace of the hair appears close to this papilla, but soon increases in length, and when the end of the hair projects from the surface, the original solid process of the epidermis becomes converted into an open pit, the lumen of which is filled by the root of the hair. Hairs differ in their mode of formation from scales in a manner analogous to that in which the teeth differ from ordinary placoid scales ; i.e. they are formed in inwardly directed projections of the epidermis instead of upon free papillae at the surface.

Nails (Kolliker, No. 298) are developed on special regions of the epidermis, known as the primitive nail beds. They are formed by the cornification of a layer of cells which makes its appearance between the horny and mucous layers of the epidermis. The distal border of the nail soon becomes free, and the further growth is effected by additions to the under side and attached extremity of the nail.

Although the nail at first arises in the interior of the epidermis, yet its position on the outer side of the mucous layer clearly indicates with which group of epidermic structures it should be classified.

Dermal skeletal structures. We have seen that in the Chordata skeletal structures, which were primitively formed of both an epidermic and dermic element, may lose the former element and be entirely developed in the dermis. Amongst the Invertebrata there are certain dermal skeletal structures which are evolved wholly independently of the epidermis. The most important of these structures are the skeletal plates of the Echinodermata.

Glands. The secretory part of the various glandular structures belonging to the skin is invariably formed from the epidermis. In Mammalia it appears that these glands are always formed as solid ingrowths of the mucous layer (Kolliker, No. 298). The ends of these ingrowths dilate to form the true glandular part of the organs, while the stalks connecting the glandular portions with the surface form the ducts. In the case of the sweat-glands the lumen of the duct becomes first established. Its formation is inaugurated by the appearance of


the cuticle, and appears first at the inner end of the duct and thence extends outwards (Ranvier, No. 311). In the sebaceous glands the first secretion is formed by a fatty modification of the whole of the central cells of the gland.

The muscular layer of the secreting part of the sweat-glands is formed, according to Ranvier (No. 311), from a modification of the deeper layer of the epidermic cells.

The Mammary Glands arise in essentially the same manner as the other glands of the skin 1 . The glands of each side are formed as a solid bud of the mucous layer of the epidermis. From this bud processes sprout out, each of which gives rise to one of the numerous glands of which the whole organ is formed. Two very distinct types in the relation of the ducts of the glands to the nipple are found (Gegenbaur, No. 313).


(304) T. H. Huxley. " Tegumentary organs." Tocld's Cyclopaedia of Anat. and Physiol.

(305) P. Z. Unna. " Histol. u. Entwick. d. Oberhaut." Archiv f. mikr. Anat. Vol. xv. 1876. FzV&also Kolliker (No. 298).

Scales of tJic Pisces.

(306) O. Her twig. " Ueber Bau u. Entwicklung d. Placoidschuppen u. d. Zahne d. Selachier." Jenaische Zeitschrift, Vol. vin. 1874.

(307) O. Hertwig. " Ueber d. Hautskelet d. Fische." Morphol. Jahrln<ch, Vol. n. 1876. (Siluroiden u. Acipenseridre.)

(308) O. Hertwig. "Ueber d. Hautskelet d. Fische (Lepidosteus u. Polypterus)." Alorph. Jahrbuch, Vol. v. 1879.


(309) Th. Studer. Die Entwick. d. Federn. Inaug. Diss. Bern, 1873.

(310) Th. Studer. "Beitrage z. Entwick. d. Feder." Zeit. f. wiss. Zool., Vol. xxx. 1878.


(311) M. S. Ranvier. " Sur la structure des glandes sudoripares." Comptes A'f/iiiits, Dec. 29, 1879.

1 For a very different view on this subject vide Creighton (No. 312).


Mammary glands.

(312) C. Creighton. "On the development of the Mamma and the Mammary function." Jour, of Anat. and Phys. , Vol. XI. 1877.

(313) C. Gegenbaur. " Bemerkungen iib. d. Milchdriisen-Papillen d. Saugethiere." Jenaische Zeit., Vol. vn. 1873.

(314) M. Huss. "Beitr. z. Entwick. d. Milchdriisen b. Menschen u. b. Wiederkauern." Jenaische Zeit., Vol. vil. 1873.

(315) C. Langer. " Ueber d. Ban u. d. Entwicklung d. Milchdriisen." Denk. d. k. Akad. Wiss. Wien, Vol. III. 1851.