The works of Francis Maitland Balfour, Volume 3 (of 4)

by Francis M. Balfour

Introduction.

Amongst the products of that part of the mesoblast which constitutes the connective tissue of the body special prominence must be given to the skeleton of the Vertebrata, from its importance in relation to numerous phylogenetic and morphological problems.

The development of the skeleton is however so large a subject that it cannot be satisfactorily dealt with except in a special treatise devoted to it; and the following description must be regarded as a mere sketch, from which detail has been as far as possible excluded.

In the lowest Chordata the sole structure present, which deserves to be called a skeleton, is the notochord. Although the notochord often persists as an important organ in the true Vertebrata, yet there are always added to it various skeletal structures developed in the mesoblast. Before entering into a systematic description of these, it will be convenient to say a few words as to the general characters of the skeleton.

Two elements, distinct both in their genesis and structure, are to be recognized in the skeleton. The one, forming the true primitive internal skeleton or endoskeleton, is imbedded within the muscles and is originally formed in cartilage. In many instances it retains a cartilaginous consistency through life, but in the majority of cases it becomes gradually ossified, and converted into true bone. Bones so formed are known as cartilage bones.

The other element is originally formed by the fusion of the ossified bases of the dermal placoid scales already described in Chapter XIV., or by the fusion of the ossified bases of teeth situated in the mucous membrane of the mouth. In both instances the plates of bone so formed may lose the teeth or spines with which they were in the first instance covered, either by absorption in the individual, or phylogenetically by their gradually ceasing to be developed. The plates of bone, which originated by the above process, become in higher types directly developed in the connective tissue beneath the skin; and gradually acquire a deeper situation, and are finally so intimately interlocked with parts of the true internal skeleton, that the two sets of elements can only be distinguished by the fact of the one set ossifying in cartilage and the other in membrane.

It seems probable that in the Reptilia, and possibly the extinct Amphibia, dermal bones have originated in the skin without the intervention of superjacent spinous structures.

In cases where a membrane bone, as the dermal ossifications are usually called, overlies a part of the cartilage, it may set up ossification in the latter, and the cartilage bone and membrane bone may become so intimately fused as to be quite inseparable. It seems probable that in cases of this kind the compound bone may in the course of further evolution entirely lose either its cartilaginous element or its membranous element; so that cases occasionally occur where the development of a bone ceases to be an absolutely safe guide to its evolution.

As to the processes which take place in the ossification of cartilage there is still much to be made out. Two processes are often distinguished, viz. (1) a process known as ectostosis, in which the ossification takes place in the perichondrium, and either simply surrounds or gradually replaces the cartilage, and (2) a process known as endostosis, where the ossification actually takes place between the cartilage cells. It seems probable however (Gegenbaur, Vrolik) that there is no sharp line to be drawn between these two processes; but that the ossification almost always starts from the perichondrium. In the higher types, as a rule, the vessels of the perichondrium extend into the cartilage, and the ossification takes place around these vessels within the cartilage; but in the lower types (Pisces, Amphibia) ossification is often entirely confined to the perichondrium; and the cartilage is simply absorbed.

The regions where ossification first sets in are known as centres of ossification; and from these centres the ossification spreads outwards. There may be one or more centres for a bone.