THE STRUCTURE 
OF 
HYALINE CARTILAGE. 
WITH DEDUCTIONS REGARDING 
THE CELL-DOCTRINE AND THE BIOPL ASSON-DOCTRINE, 
BEING THE FIRST CHAPTER OF “CONTRIBUTIONS TO THE HIS- 
TOLOGY OF THE CARTILAGES OF THE LARYNX.” S 
By LOUIS ELSBERG, A.M., M.D., 
PROF. OF LARYNGOLOGY AND DISEASES OF THE THROAT IN DARTMOUTH MEDICAL COLLEGE. 
Reprinted from the Archives of Laryngology, Vol. ii, No. 4, October, 
1881 and Vol. iii, No. 1, January, 1882. Plate of the Thyroid Cartilage of Adult. Longitudinal Section, x 100. 
A. —Perichondrium toward the mucous membrane. 
B. —Perichondrium toward the skin. 
F.—Fibrous portion of cartilage in the centre. 
H.—Hyaline portion, on either side, near the perichondrium. 
Fibrous Portion of Thyroid Cartilage, x 600. 
C.—Cartilage corpuscle, surrounded by a dense basis-substance.. 
G.—Granules of living matter. 
B.—Capillary blood-vessel. 
V.—Small vein. 
FIG. 2. 
FIG. I. 
ELSBERG’S CONTRIBUTIONS TO THE HISTOLOGY OF THE CARTILAGES OF THE LARYNX.—(See page 335.) 
Thyroid Cartilage of Adult, kept in strong Alcohol. Horizontal Section, x 1200 
C.—Shrivelled cartilage corpuscle. 
O.—Longitudinal offshoots. 
B.—Reticulum in basis-substance. 
G.—Granules of living matter. 
Thyroid Cartilage of Adult. Sagittal Section, x 1200. 
C.—Cartilage corpuscles. 
B.—Indistinctly reticular hyaline basis-substance. 
F.—Fibrous basis-substance. 
FIG. 4. 
FIG. 3. 
Thyroid Cartilage of Adult. Horizontal Section, x 1200. 
C.—Cartilage corpuscle. 
B.—Hyaline basis-substance. 
G.—Granules of living matter. 
Thyroid Cartilage of Adult. Horizontal Section, x 600. 
C.— Cartilage corpuscles. 
F. — Fibrous portion of cartilage. 
G. Granules of living matter. 
FIG. 6. 
FIG. 5. CONTRIBUTIONS TO THE NORMAL AND PATH- 
OLOGICAL HISTOLOGY OF THE CAR- 
TILAGES OF THE LARYNX. 
By LOUIS ELSBERG, M.D., 
NEW YORK. 
CHAPTER I.—ESSENTIAL STRUCTURE OF HYALINE 
CARTILAGE* 
Section I.—Historical. 
FROM the earliest time of histology to the present, 
true cartilage, such as the thyroid cartilage, has been 
looked upon as one of the simplest tissues. To distinguish 
it from other kinds of cartilage, in which either a fibrous or 
a reticular aspect has been recognized, it is called hyaline, 
i. e., resembling glass. The description of its structure, by 
Meckauer in 1836,1 is essentially as that by Klein in 1880,* 
viz., that it consists of a firm homogeneous basis-substance, 
in which are imbedded numerous small cartilage-corpuscles. 
Meckauer wrote before the cell-doctrine, which has exercised 
so powerful an influence upon the medical mind, had been 
thought of. Indeed, that doctrine itself, as its founder, 
Schwann,3 has recorded, was based to a large extent upon 
* Read before the American Laryngological Association, session 1881. 
1 De penitiori cartilaginum structura symbolae. Diss. anat.-phys., auctore 
M. Meckauer, M.D. Breslau : Schultz & Co., 1836, Tab. 4, pp. 16. 
3 Atlas of Histology. London : Smith, Elder & Co., 1880, p. 48. 
3 Mikroskopische Untersuchungen tiber die Uebereinstimmung in der Structur 
und dem Wachsthume der Thiere und Pflanzen., von Dr. Th. Schwann. Ber- 
lin : G. E. Reimer, 1839, pp. 270. Microscopical researches into the accord- 
ance in the structure and growth of animals and plants. Translated by Henry 
Smith. London : Sydenham Society, 1847. Introduction. 
Reprinted from the Archives of Laryngology, Vol. ii, No. 4, Oct., 1881. 2 
Louis Elsberg. 
investigations of the constitution of cartilage. After J. 
Muller had described cartilage-corpuscles that were hollow, 
and Gurlt had spoken of some as vesicles, when Schwann 
had succeeded, as he thought, “ in actually observing the 
proper wall of the cartilage-corpuscles, first in the branchial 
cartilages of the frog’s larvae, and subsequently also in the 
fish,” he was led by these and other researches to conjecture 
“ that the cellular formation might be a widely extended, 
perhaps a universal principle for the formation of organic 
substances.” 
Schwann considered that the cartilage-corpuscles, or car- 
tilage-cells as they were thenceforth called, are imbedded in 
a matrix which is capable of producing the cells, and which 
he therefore called cytoblastema. Goodsir, Naegeli, and 
finally Virchow advanced the histology of cartilage in so 
far as they claimed that the cartilage-cells can not possibly 
arise from the matrix or intercellular substance. Even 
Virchow adhered, however, to the idea of Schwann, that 
the cartilage-cell is a vesicle filled with a more or less 
transparent fluid, in which is suspended the nucleus; and, 
although he was aware of the life of the cell in general, 
nothing was suggested by him as to the life of cartilage. 
It is true Donders and H. Meyer had observed that the cells 
of hyaline cartilage were capable of proliferation1; neverthe- 
less the idea became prevalent, more perhaps from implica- 
tion, because on account of the absence of blood-vessels 
it was believed not liable to inflammation, than from any 
direct statement to that effect, that cartilage was devoid 
of life. The vitality of cartilage-corpuscles was made 
clearly probable by the observation of the effect of elec- 
trical shocks upon them, by Heidenhain,2 and by Rollett,8 
and the investigations of Reitz,4 Boehm,6 Hutob,* and Bub- 
1 (Mueller’s) Archiv fur Anatomie, 1846. 
a Studien aus dem physiologischen Institut zu Breslau, ii Heft, 1863. 
3 Strieker’s Handbuchder Lehre von denGeweben : Article “ Knorpelgewebe,” 
1868. 
* Sitzungsber. der K. K. Akademie der Wissensch. in Wien., Bd. 55, 1867. 
6 Beitrage zur normalen und pathologischen Anatomie der Gelenke. Inaug.- 
Dissertation, Wiirzburg, 1868. 
8 “ Untersuchungen Uber Knorpelentztindung.” Wiener. tried. Jahrbiicher, 
1871, p. 399. Histology of the Cartilages of the Larynx. 
3 
noff,1—investigations which, except Boehm’s, were made 
under Strieker; it was proved positively by Heitzmann in 
1873.2 
With the question whether or not the so-called cartilage- 
“ cell ” is alive, another question arose, viz., how can so 
isolated a corpuscle (imbedded in a firm “ intercellular ” sub- 
stance) obtain nutrition ? It was assumed that the nourish- 
ing liquid reaches the corpuscle either by diffusion or else 
through canals, or clefts, or fissures in the homogeneous 
basis-substance. The idea of the existence of juice-channels 
originated with Von Recklinghausen. He found in silver- 
stained preparations of the cornea, communicating colorless 
spaces on a dark background, and believing that the cornea 
consisted of fibrillary tissue knit together by a cement-sub- 
stance, he thought that this cement-substance was tunneled 
by a system of communicating-canals, “ Saft-Kandlcheii,” 
and that it is this system of canals which is not stained by 
silver. Innumerable investigations, under all sorts of cir- 
cumstances, have been undertaken to settle satisfactorily 
whether preformed juice-channels exist in cartilage, or 
whether juices can be imbibed without such. In lower ani- 
mals corresponding canals had long been reported to be 
found, by Queckett3 and by Bergmann4 in cephalopodes, and 
by Leydig5 in various fishes; and certain pathological obser- 
vations by Virchow',8 Zahn,7 Cornil and Ranvier,8 and Rind- 
fleisch,9 as well as senile changes studied by Weichselbaum,10 
1 Beitrage zur Kenntniss der Structur des Knorpels. Sitzungsber. der K. K. 
Akad. d. Wiss. in Wien., Bd. 57, 1868. 
3 “ Das Verhaltnisszwischen Protoplasma und Grundsubstanzim Thierkbrper. ” 
Sitzungsber. d. K. K. Akad. d. Wien. Wien., Bd. 67, 1873, and Wien. med. 
Jahrbiicher, 1873. 
3 Catalogue of the historical series in the museum of the Royal College of Sur- 
geons, 1850, vol. i, p. 102. 
4 Disquisitiones microscopicse de cartilaginibus in specie hyalinicis. Inaug.- 
Dissert., Dorpat, 1850. 
6 “ Zur Anatomie und Histologie der Chimsera monstrosa.” Mueller's Archiv, 
1851, p. 242. 
6 “ Ein Fall allgemeiner Ochronose der Knorpel und knorpelahnlichen Theile.” 
Virchow's Archiv, xxxvii, 1866, p. 212. 
7 “ Uber Pigmentinfiltration des Knorpels.” Ibid., lxxii, 1878. 
8 Manuel d’ histologie pathologique, Paris, 1869, p. 427. 
* Lehrbuch der pathologischen Gewebelehre. Leipzig, 1878, p. 553. 
10 Sitzungsber. der K. K. Akademie d. Wiss. in Wien., Bd. 75, 1877. 4 
Louis Elsberg. 
seemed to point to their presence in man. Pigment particles 
were introduced into the circulation in the hope of discover- 
ing the manner in which they penetrate the tissue of carti- 
lage, by Gerlach,1 Maas,2 Arnold,3 and Nykamp and Treub4; 
Kiittner, with the same end in view, introduced solutions 
into the trachea and examined the bronchial and tracheal 
cartilages6; and Henoque," Budge,7 Tizzoni,8 and others, 
forcibly injected liquids as well as solid particles into the 
tissues. The results of these experiments, and of examina- 
tions with various reagents, are contradictory of each other: 
For instance, while Bubnoff,8 Hertwig,10 Henoque,11 Loewe,12 
Thin,13 Ewetzky,14 Petrone,16 Budge,18 Nykamp,17 Furbringer,18 
and a number of others consider the existence of canals in 
the basis-substance of cartilage proved by their experiments 
and treatment of their preparations with silver nitrate, gold 
chloride, hyperosmic acid, chromic acid, ammonia bichro- 
I Ueber das Verhalten des indigschwefelsauren Natrons im Knorpelgewebe 
lebender Tliiere. Erlangen, 1876. 
3 “ Ueber das Wachsthum und die Regeneration der Rbhrenknochen.” Ar- 
chiv fiir klinische Chirurgie, xx, 1877. 
3 “ Die Abscheidung des indigschwefelsauren Natron im Knorpelgewebe.” 
Virchow's Archiv, lxxiii, 1878. 
4 “ Beitrag zur Kenntniss der Structur des Knorpels.” Archiv fiir mikro- 
skop. Analomie, xiv, 1877. 
6 “ Die Abscheidung des indigschwefelsauren Natron in den Geweben der 
Lunge.” Centralblatt f. d. med. IViss., 1875, No. 42, p. 268. 
8 “ Structure des cartilages.” Gazette mddicale, 1873, p. 589 ; p. 617. 
7 “ Die Saftbahnen im hyalinen Knorpel.” Archiv fiir mikroskop. Anatomie, 
xiv, 1877 ; xvi, 1879. 
8 “ Sulla istologica normale e patologica delle cartilagini ialini.” Archivioper 
le Scienze Mediche, ii, 1877. 
9 Loc. cit. 
10 “ Ueber die Entwickelung und den Bau des elastischen Gewebes im Netz- 
knorpel.” Archiv fur mikroskop. Anatomie, ix, 1873, p. 80. 
II Loc. cit. 
13 “ Ueber eine eigenthttmliche Zeichnung im Hyalinknorpel.” Wiener med. 
Jahrbiicher, 1874. 
13 “ On the structure of hyaline cartilage.” Quarterly Journal of Microscopical 
Science, vol. xvi, 1876. 
14 “ Entziindungsversuche am Knorpel,” Vorlaufige Mittheilung, Centralblatt 
f. d. med. Wiss., 1875, No. 16; Untersuchungen aus dem path.-anat. Institut 
zu Zurich, iii Heft, 1875. 
16 Sulla struttura normale e patologica delle cartilagine e degli epitelii. Na- 
poli, 1876. 
18 Loc. cit. 17 Loc. cit. 
18 “ Ueber das Gewebe des Kopfknorpels der Cephalopoden.” Morpholog. 
Jahrbiicher, iii, 1877, p. 453. Histology of the Cartilages of the Larynx. 
5 
mate, etc., etc., investigations by exactly the same means have 
convinced Sokolow,1 Retzius,2 Colomiatti,3 Bruckner,4 Toldt," 
Genzmer,6 Gerlach,7 Tillmanns,8 Tizzoni,9 and others, of just 
the contrary ; and there is a third party which believes with 
Arnold10 that the basis-substance is made up of fibrillae, that 
there are delicate fissures between the fibrils, that these fis- 
sures penetrate the capsule, and that “ the nutrient material 
passes through these interfibrillar and intracapsular fissures 
into the pericellular space.” Flesch, the latest writer on 
the subject, adds 11 that these fissures need not necessarily 
be, and in fact are not, empty, but that they are occupied 
by the interfibrillar cement-substance, which, being of a 
“ viscous-soft ” (zahweich) material, permits the imbibition 
and conveyance of the nutrient liquid. 
It is claimed that hyaline basis-substance consists of fine 
fibrils so closely held together by a cement-substance that 
the mass appears to be homogeneous. This idea, though 
not entirely novel, as the older anatomists seem to have had 
it,12 has been brought forward by Tillmanns, and is doubtless 
original with him.13 It is said that the interfibrillar cement- 
substance can be dissolved out by certain reagents and then 
1 Ueber den Bau des Nasenknorpels,” etc., ref. Canstatt’s Jahresbericht. 
1870, p. 24. 
2 “ Bitrag till Kannedomen um brusknafnaden.” Nord. med. Arkiv., iv, 
1872. 
3 “ Sulla struttura delle cartilagini ialini e fibroelastica reticolata.” Gazetta 
Cliniche di Torino, 1873, No. 32 ; Rivista Clinica di Bologna, 1874, No. 5 ; 
Giornale della Acad, di Torino, 1876. 
4 “ Uber Eiterbildung im hyalinen Knorpel.” Inaug.-Dissert., Dorpat, 1873. 
6 Lehrbuch der Gewebelehre. Stuttgart, 1874, p. 143. 
6 “ Ueber die Reaction des hyalinen Knorpels,” etc. Virchow's Archiv, 
Ixvii, 1875 ; Centralblatt f. Chirurgie, 1875, No. 146. 
7 Loc. cit. 
8 “ Beitrage zur Histologie der Gelenke.” Archiv fur mikroskop. Anatomic, 
x, 1874, PP- 354. 435- 
9 Loc. cit. 10 Loc. cit. 
11 Untersuchungen liber die Grundsubstanz des hyalinen Knorpels. Wurz- 
burg : A. Stuber, i860. 
12 See : Wm. Hunter “ On the structure and diseases of articular cartilages,” 
Philosoph. Transactions, vol. xlii, p. 514, London, 1742-43; M. de Lasone, 
“ Second memoire sur 1’ organization des os,” Mem. de 1’ Academie Roy. des 
Sciences, Tome 69, Paris, 1752 ; more recently also : Hoppe, Virchow’s Archiv, 
v, p. 175. 
13 Loc. at., p. 401 ; and “ Ueber die fibrillare Structur des hyalinen Knorpels.” 
Archiv f. Anatomic u. Physiologic, Anat. Abth., 1877, p. 9. 6 
Louis Els berg. 
the fibrillation seen under the microscope. According to 
the varying arrangement and interrelation of the fibrillae, 
Tillmanns speaks of three types of cartilage tissue, viz., 
parallel-fibery, netform, and lamellous. No doubt he saw 
under the microscope appearances which underlie the dis- 
tinction which he thus made, but, unfortunately, he misin- 
terpreted these appearances. Nevertheless, he has had 
followers. Thus Baber reported 1 that, having undertaken 
to test the accuracy of Tillmanns’ assertions, and not suc- 
ceeding in finding the fibrillation, although he had followed 
Tillmanns’ method of maceration, he accidentally made mo- 
mentary pressure on the glass cover, and thereupon obtained 
satisfactory proof of the fibrillar constitution of the basis- 
substance. Reeves2 has also convinced himself of the exist- 
ence of normal fibrillation in human cartilage. Ziegler seems 
to have done the same 3; and Flesch regards it as a matter 
beyond question. He speaks of it as “ generally known and 
most easily demonstrable.” 4 Furthermore, he thinks that 
some portions, or perhaps layers, of the basis-substance 
are more compact than others, and that this may also 
account for the facility of cleavage in determinate direc- 
tions. 
Leidy insisted5 that the basis-substance of hyaline car- 
tilage has a peculiar filamentous structure, but his interpre- 
tation, that the granular filaments run simply parallel to each 
other, does not cover the truth and has not attracted any 
attention. With the exception of Leidy, however, no 
one, until nine years ago, seems to have questioned the 
homogeneousness of the mass of basis-substance in which 
the separate corpuscles were supposed to be imbedded. In 
1872, Heitzmann* first proved the presence of a network 
structure in the basis-substance ; one year later he discov- 
1 “ On the structure of hyaline cartilage.” Journal of Anatomy and Physi- 
ology, vol. x, part i, October, 1875. 
2 “ On the structure of the matrix of human articular cartilage.” British 
Medical Journal, Nov. II, 1876, p. 616. 
3Bericht der 50. Naturforsch. Versammlung zu Mtinchen, 1877. 
4 Loc. cit., p.,74- 
6 Proceedings of the Academy of Natural Sciences of Philadelphia, vol. iv, 
No. vi, 1848 ; and American Journal of Medical Sciences, April, 1849, p. 282. 
6 Wiener Medizin. Jahrbiicher, Heft iv, 1872. Histology of the Cartilages of the Larynx. 
7 
ered the structure of so-called protoplasm to be reticular, 
and claimed that both the reticulum of the protoplasm and 
that of the basis-substance are identical in nature; viz.: 
that both consist of the living matter proper. Heitzmann 
says that examination, by means of an immersion lens 
No. 10, of a thin section of fresh articular cartilage, placed 
in a one-half-per-cent, solution of common table-salt reveals 
details heretofore overlooked, viz.: The bodies of the cells 
appear finely granular, bounded by a somewhat denser 
layer. The contour of a cartilage cell being accurately in 
focus, there appears between it and the basis-substance a 
light, very narrow rim, which is traversed by numerous ex- 
tremely delicate, radiating, grayish thorns or streaks. All 
these thorns are conical, the broad base emanating from the 
body of the cell and the thin point directed toward the 
basis-substance. Wherever two cells lie close together, the 
light rim between them is pierced by grayish threads. 
When in a cell the nucleus is distinctly seen, a narrow light 
rim is found to surround it, which on being sharply focused, 
also shows radiating thorns, the basis of which emanate 
from the nucleus and the points of which blend with the 
protoplasm of the cell. On carefully examining the basis- 
substance, a very delicate, as if granular, configuration is 
recognizable, dark fields alternating with light ones, and in 
some places the impression is given that the light fields 
form ramifications, or even a delicate network. Heitzmann 
also described and figured specimens stained with silver 
nitrate and gold chloride, and announced the following con- 
clusions: viz.: “The bodies of cartilage cells have radiating 
offshoots. These offshoots form a delicate granular reticu- 
lum in the basis-substance. At the points of junction of 
hyaline cartilage with fibrous cartilage and with periosteum, 
the offshoots are very large and broad. They connect 
neighboring cells either directly or else indirectly through 
intervening delicate offshoots.” Somewhat similar appear- 
ances had previously been more or less vaguely described, 
but not properly interpreted or appreciated, Remak,1 
J “ Ueber die Entstehung des Bindegewebes und des Knorpels.” Archiv fur 
Anatomie, 1852, p. 63 et seq. 8 
Louis Els berg. 
by Heidenhain,1 by Broder,2 by Fromann,8 and possibly by 
others. 
After Heitzmann, Hertwig4 observed processes of living 
matter penetrate the basis-substance of reticular cartilage; 
and Colomiatti stated 6 that he had repeated the investiga- 
tions of Heitzmann, but had failed to find cell offshoots in 
hyaline cartilage either after treatment with gold or silver 
or in vivo, although he had seen cartilage-cell offshoots in 
other than hyaline cartilage. 
I have had the opportunity to repeat Heitzmann’s inves- 
tigations under his own eye and with his assistance, but the 
results as to their correctness at which I arrived, were, to 
the best of my belief, uninfluenced by him. I reported in 
1875 6 that I had seen the network structure in the corpus- 
cles of hyaline cartilage, in the nucleus and in the basis- 
substance, exactly as Heitzmann had described it two years 
previously.7 “ If the nucleus of the cartilage corpuscle be 
visible, it appears either homogeneous or composed of a 
dense meshwork of living matter. From its periphery pro- 
ceed fine conical thorns, which lead to a meshwork pervading 
the whole corpuscle, the threads of which form, at the 
point of intersection, thickenings, granules, or small clumps 
of living matter. In the lighter-looking narrow seam, ex- 
isting between the corpuscles and the surrounding matrix, 
we may also recognize fine threads which go from the pe- 
riphery of the corpuscle and are lost to the view in the ma- 
trix. On examining such a fresh preparation upon the 
heated stage, we recognize, at a temperature of 86° to 950 
F., a continual but very slow change in the living matter of 
those cartilage corpuscles which distinctly show the net- 
work ; the points of intersection of threads move nearer 
1 Loc. cit. 
3 Ein Beitrag zur Histologie des Knorpels. Dissert., Zurich, 1865. 
3 Untersuchungen Uber die normale und pathologische Anatomiedes Rttcken- 
markes. II Theil. Jena, 1867, pp. 29, 30. 
4 Loc. cit. 6 Loc. cit. 
® Transactions of the American Medical Association, vol. xxvi, 1875, pp. 
163 and 164. 
7 Untersuchungen uber das Protoplasma. II. Das Verhaltniss zwischen Pro- 
toplasma und Grundsubstanz im Thierkbrper. Sitzungber. d. K. K. Akad. d. 
Wiss. in Wien, Ixvii, May, 1873. Histology of the Cartilages of the Larynx. 
9 
together or go further apart ; sometimes a few granules lying 
close to each other unite into one little lump, so that the 
threads between them disappear, then the latter reappear, 
lengthen and shorten ; and this change in the form of the 
interior network continues for some little time, and without 
any perceptible influence upon the form of the corpuscle as 
a whole. 
“Careful examination of the matrix reveals throughout 
its whole extent the existence of a very delicate, more or 
less distinct network of living matter; and in many instances 
the connection of the threads proceeding from the corpus- 
cles with this delicate network can be traced. 
“To show the structure of both the cartilage corpuscles 
and the matrix still more plainly, we may resort to the 
method of tinction of preparations by nitrate of silver and 
chloride of gold, as well as the examination of cartilage 
during normal calcification, and in its inflamed conditions. 
“ It is well known that chloride of gold stains living matter 
dark violet, while nitrate of silver acts upon the matrix, and 
by darkening it, makes the living matter appear of a light 
color or colorless. The appearances obtained thus comple- 
ment each other; and the network proceeding from the 
corpuscles and ramifying all through the matrix, is seen, 
with the same magnifying power, as constituted by violet 
threads and granules in the first case, and by white processes 
or empty spaces in the second case. [Drawings illustrating 
both these appearances accompany my article in the 
American Transactions.] As the deposition of lime salts 
takes place only in the matrix, the living matter itself re- 
maining free, careful examination during such depositions, 
especially in cases of artificially produced inflammation, also 
brings to view the fine network of living matter traversing 
the matrix.” 
In January, 1876, Thin’s memoir was published,1 in which 
he reported that, in particular preparations, he had seen “fine 
glistening fibres enter the cartilage substance, into which, 
however, he has not been able to follow them.” Again : 
“The ordinary granular protoplasmic cells of hyaline carti- 
1 It is dated August, 1875. loc. cit. 10 
Louis Els berg. 
lage are analogous, according to the views of the author, to 
the stellate cells of the cornea and connective tissue gener- 
ally.” Thin obtained by silver staining appearances similar 
to Heitzmann’s, but unfortunately misinterpreted them. 
He says: “ Heitzmann believes that the appearances which 
he has reproduced are those of a cell and its protoplasmic 
processes. The author interprets the appearances shown in 
Heitzmann’s own drawings as representing stellate spaces, 
and sees nothing in them that he can interpret as cell pro- 
cesses; being thus at one with him as regards the fact 
observed, but differing from him in regard to its interpreta- 
tion.” 1 
In 1879 Spina reviewed the subject.2 He accorded to 
Heitzmann the merit of the discovery, but as in the inter- 
vening seven years I alone had publicly corroborated it, and 
he was not aware of that corroboration, he thought that 
“the existence of cells with solid offshoots in genuine hya- 
line cartilage is not definitely proved,” and undertook to 
settle the question. After many fruitless attempts he found 
out a method of examination “ by which ramifying cells in 
hyaline cartilage can be demonstrated not only with ease, 
but also with certainty.” The method and the results, as 
he has described them, are as follows : “ The cartilage, best 
the articular ends of bones, is placed into alcohol for three 
or four days; then the sections are made and the examina- 
tion is conducted in alcohol. From such specimens positive 
proof is obtained that the cells of hyaline cartilage have 
solid offshoots. These offshoots emanate mostly from the 
body of the shrivelled cells, penetrate the basis-substance, and 
inosculate with offshoots of other cells. Their number and 
thickness are subject to numerous variations. * * * The 
cell offshoots do not, as a rule, ramify. * * * Examina- 
tion with powerful immersion lenses (Hartnack, No. 15) 
teaches positively that the cell offshoots not only pierce the 
capsule, but that the capsule extends also to the offshoots 
themselves, so that at their origin they are surrounded like 
the cell body by a wall. * * * Upon adding a drop of 
1 Loc. cit., p. 22. 
2 “ Ueber die Saftbahnen des hyalinen Knorpels.” Sitzungsber. der K. K. 
Akad. d. Wiss. in Wien, lxxx, Abth. iii, Nov., 1879. Histology of the Cartilages of the Larynx. 
11 
glycerine to the alcohol specimen, or on staining it after 
one of the usual methods, the cell offshoots disappear more 
or less rapidly; hence, it is clear that the hyaline, structure- 
less aspect of the cartilage basis-substance, is really due to 
the methods of preparation hitherto in use, while, when ex- 
amined in alcohol, as above described, the cell offshoots in- 
variably become visible.” He added that he has succeeded 
a few times in seeing—faintly only, it is true—the same struc- 
ture in living hyaline cartilages. On incorporating, for a 
sufficient length of time, carmine into the body of frogs, 
Spina found cartilage corpuscles of which the nuclei, the 
body, and the offshoots had taken in some of the coloring 
matter. As the offshoots disappeared and the carmine 
granules seemed to lie in the hyaline basis-substance when 
a drop of glycerine was added, it is easy to see how previ- 
ous investigators came to be misled into supposing the 
coloring matter to have passed into the hyaline substance, 
and into interfibrillar fissures. With excessive caution, 
Spina adds: “Whether they (the coloring particles) can also 
move along outside of the cell offshoots has not yet been 
proved.” 
In the same year Prudden,1 and, in 1880, Flesch,2 also 
described cilia-like processes of cartilage corpuscles; and 
the latter admitted that in exceptional cases he had suc- 
ceeded in tracing them more or less distinctly into the 
basis-substance. 
Section II.—Personal Investigation. 
I. Having, as already detailed, familiarized myself with 
the investigation of cartilage tissue, I entered upon the ex- 
amination of the laryngeal cartilages. Of the many speci- 
mens examined, I shall now describe a few of the thyroid. 
Longitudinal sections through the lateral plates of the thy- 
roid cartilage of a man of about 25 years, hardened in chro- 
mic acid and stained with an ammoniacal carmine solution, 
exhibit with low powers of the microscope (150 to 200 
1 “ Beobachtungen am lebenden Knorpel.” Virchow's Archiv, lxxv, 1879, 
p. 185. 
2 Loc. cit., pp. 59-63. Louis Elsberg. 
12 
diam.) the following: The cartilage corpuscles, either single, 
in pairs, or in groups of from three to six, or even more, are 
imbedded in a basis-substance which, for the most part, is 
homogeneous-looking or indistinctly granular, but in some 
portions finely striated. The homogeneous or indistinctly 
granular-looking basis-substance is that which bears the name 
hyaline basis-substance; the striated is termed fibrous, al- 
though actual fibrillae appear only on the edges of the speci- 
men, or when the tissue is torn and mutilated. The fibrous 
basis-substance is intermixed, without any regularity, with 
the hyaline, and usually sharply separated from it. Not infre- 
quently a number of cartilage corpuscles, or groups of carti- 
lage corpuscles, are surrounded by fibrous basis-substance, 
the striations of which run, as a rule, in a sagittal direction, i.e., 
vertical to the surface. Within the fibrous basis-substance 
the cartilage corpuscles are at most points sparsely scattered 
or absent; here and there, however, they are more numer- 
ous, in rows or elongated, corresponding to the direction of 
the striations. It also occurs that striated portions of the 
basis-substance contain very minute globular or oblong cor- 
puscles, sometimes to such an extent that the striated struc- 
ture is concealed by the large number of these corpuscles. 
Fig. i exhibits the appearance of the constituent parts, 
with an amplification of ioo diam. The fibrous portion is 
seen to occupy the centre of a longitudinal section of one 
of the plates of the thyroid cartilage. This is not regularly 
the case in every cut, and was exceptionally well marked in 
the section from which the drawing was made. In some 
sections the fibrous cartilage is altogether absent, but every 
laryngeal cartilage contains some fibrous mixed with hya 
line portions. A, perichondrium toward the mucous mem- 
brane. B, perichondrium toward the skin. F, fibrous por- 
tion of cartilage. H, hyaline portion of cartilage. 
Under higher magnifying powers (500 to 600 diam.) single 
cartilage corpuscles exhibit features, frequently before de- 
scribed, with coarsely granular nuclei. Around the nucleus 
finer granules are visible. At the periphery of the cartilage 
corpuscle there are several strata of higher refracting power, 
especially the zone nearest the basis-substance, which, as a Histology of the Cartilages of the Larynx. 
13 
rule, appears very shining and is what is termed the capsule 
of the cartilage corpuscle. Not infrequently the cartilage 
corpuscle is very indistinct, being but slightly more granu- 
lar than the surrounding basis-substance; then almost 
nothing but the nucleus marks its presence and its place. 
In twin formations of cartilage corpuscles, which are often 
met with, the zone of division between the two corpuscles 
is identical with that surrounding both, in the shape of a 
capsule. Of the same nature are the zones of division that 
are seen in clusters of cartilage corpuscles. 
The so-called hyaline basis-substance throughout its 
whole extent now appears finely granular; as a rule, the 
granulation is more distinct midway between the corpuscles 
than in their immediate vicinity. The fibrous portions of 
the basis-substance are seen to be made up of extremely 
minute spindles, which, by being grouped longitudinally, 
produce the aspect of striation. The spindles or fibres are 
separated from each other by light rims, and both the 
spindles and the rims look finely granular. Between the 
spindles may often be seen small globular bodies, sometimes 
scattered, sometimes in clusters, of which the size and shape 
greatly vary, reaching occasionally the size and shape of a 
regular cartilage corpuscle. In some striated fields blood- 
vessels, both arterial and capillary, can be seen ; the former 
with the characteristic muscle-coat, the latter with the en- 
dothelial wall, besides holding red blood corpuscles in their 
calibres. 
Fig. 2, representing the fibrous portion of thyroid carti- 
lage amplified 600 diam., shows C, a cartilage corpuscle sur- 
rounded by a dense basis-substance; £, granules of living 
matter in a dense fibrous structure ; B, a capillary blood- 
vessel ; and V, a small vein. 
The highest powers of the microscope (1000 to 1200 
diam.) reveal the reticular structure of cartilage corpuscles, 
as it is known since 1873. All granules within the nucleus 
and all granules within the corpuscle are uninterruptedly 
connected by delicate threads. The intranuclear network 
is connected with the corpuscular reticulum by radiating 
conical spokes traversing the light rim around the nucleus; 14 
Louis Els berg. 
and, at the periphery of the corpuscle, similar conical spokes 
pierce a narrow light rim and enter the basis-substance, in 
which, especially in the highly refracting zone, termed cap- 
sule, they are usually lost to sight. Cartilage corpuscles 
even which have become so pale as to leave only a dim trace 
of their former contour visible, still exhibit more or less dis- 
tinct traces of the reticular structure. 
The same structure may be seen throughout the so-called 
hyaline basis-substance—more distinct in the middle of the 
space between the corpuscles than immediately around the 
corpuscles themselves. The fibrous portion of the basis- 
substance has also a reticular structure. The bodies of the 
slender spindles show a network without the application of 
any reagent, and the light rims between the spindles are tra- 
versed by delicate threads running in a vertical direction to 
the longitudinal diameter of the spindles. All granules and 
lumps scattered through the fibrous basis-substance are sur- 
rounded by light rims, which are pierced by conical spokes 
inosculating with the reticulum of the neighboring spindles. 
The reticular structure of cartilage corpuscles, C C, twin- 
formations ; the indistinct reticulum in hyaline basis-sub- 
stance, B; and a number of small granules of living matter 
in fibrous basis-substance, F, the spindles and granules being 
connected by fine threads, are exhibited in fig. 3 which 
represents a sagittal section of the thyroid cartilage of an 
adult amplified 1200 diam. 
I have treated sections of the same cartilage after they 
had for several days been washed out with distilled water, 
with a one-half-per-cent, solution of gold chloride, where- 
upon they assumed a dark purple color, and showed all the 
features described, somewhat more distinctly than simple 
carmine preparations. 1 deem their detailed description 
unnecessary. 
II. When I became acquainted with Spina’s researches, 
cited in my historical sketch, I deemed it of importance to 
repeat the examination according to his method. I there- 
fore placed a larynx immediately after removal from the 
body of a girl, aged 24 years, into strong alcohol, and after 
four days made thin sections from the thyroid cartilage in Histology of the Cartilages of the Larynx. 
15 
a horizontal direction, transferred them in alcohol to the 
slide, and examined them with both low and high powers, 
adding from time to time a drop of strong alcohol to pre- 
vent the specimen from drying. The appearance presented 
by such a specimen is truly surprising. As a matter of 
course, the cartilage corpuscles are shrivelled up so that 
more or less space is left between their jagged periphery and 
the border of the basis-substance. With an amplification 
of 500 diam., the basis-substance is seen pierced by light fila- 
ments, which, in many instances, can be traced through the 
intervening space into the body of the cartilage corpuscle. 
Most of these filaments radiate around the corpuscle, and 
immediately after penetrating the basis-substance, diverge 
and form a reticulum throughout its extent. Cartilage 
corpuscles located near each other, are directly connected 
by non-ramifying and occasionally by ramifying offshoots, 
or by bundles of such offshoots of a more or less parallel 
course. The reticulum in the basis substance is either radi- 
ating or irregularly arranged around the corpuscle. Con- 
trary to the assertion of Spina, the filaments or offshoots 
do, as a rule, ramify, except those that directly connect the 
neighboring corpuscles. Sometimes thick bundles of off- 
shoots emanate from opposite poles of the corpuscles, while 
intervening portions of the periphery are almost devoid of 
offshoots. Toward the periphery of the thyroid cartilage, 
—where, as is well known, the cartilage corpuscles elongate, 
becoming smaller and spindle-shaped and more or less par- 
allel to each other,—the offshoots are given off rectangular- 
ly to the axis of the corpuscles. 
High magnifying powers, immersion lenses No. 10 and 
No. 12, conclusively prove the connection of the offshoots 
with the cartilage corpuscles. Portions of the basis-sub- 
stance which, with lower powers, looked only granular, now 
show a delicate reticulum, which, even when coarser off- 
shoots are wanting, is connected with the cartilage corpuscle 
through delicate, and more or less conical, offshoots from 
the surface of the corpuscle. 
The light interstices between the fibres of striated basis- 
substance, are also traversed by delicate grayish thorns. 16 
Louis Elsberg. 
Such thorns are visible even in the perichondrium. Through 
the fibrous bundles of the perichondrium run in a nearly 
rectangular direction, delicate light streaks, while the inter- 
stices between the bundles and the spaces left between the 
corpuscular elements and the bundles, exhibit delicate coni- 
cal grayish threads, the direction of which corresponds to 
these light streaks. 
The highest powers of the microscope disclosed in one of 
the specimens examined another feature in the hyaline 
basis-substance, viz., the presence ot a number of granules 
or minute lumps of varying shape, some interwoven with the 
direct offshoots of the corpuscles, and some with the threads 
forming the finer network of the basis-substance. They ap- 
peared to be thickened points of intersection, knots or nodes, 
composed of the same material as the offshoots and threads 
themselves. They were unquestionably granules of living 
matter. I found their greatest development in a case ex- 
amined without Spina’s method, a case which I shall de- 
scribe presently. Fig. 4 shows a horizontal section of a 
thyroid cartilage, which was hardened in alcohol and exam- 
ined by the method of Spina, with an amplification of 1200 
diam. C, shrivelled cartilage corpuscle; O, longitudinal off- 
shoots connecting cartilage corpuscles ; R, reticulum in the 
hyaline basis-substance ; G, granules of living matter, which 
are seen to be part and parcel of the reticulum. 
III. The observation which I am now about to record, 
was made in specimens of the thyroid cartilage removed 
from the body of a rather stout man, 48 years old. After 
having been hardened in chromic acid solution, without any 
other reagent, they exhibited formations in the basis-sub- 
stance which, so far as I am aware, have never before been 
described. I have alluded to them as found in one of the 
specimens examined, with the highest powers of the micro- 
scope, by the alcohol method of Spina. These formations 
are shown in fig. 5, with an amplification of 600, and in 
fig. 6, with an amplification of 1200 diameters. 
As to the cartilage corpuscles in these specimens, many of 
them were larger and more coarsely granular than are com- 
monly observed : otherwise, their characters and the arrange- Histology of the Cartilages of the Larynx. 
17 
merits of the basis-substance, both so-called hyaline and 
fibrous, were like those described before. The intranuclear, 
intracorpuscular, and intercorpuscular networks were with 
high powers well shown. 
The very remarkable feature was that with quite low 
power the basis-substance was seen to be speckled and stud- 
ded with granules or lumps varying from that of a point at 
the limit of the visible to that approaching the dimensions 
of a regular cartilage corpuscle. Of course, no one must for 
a moment think of any thing like the pathological condi- 
tions that have been described either as granular degenera- 
tions of the cartilage basis-substance, or as incrustations of 
the corpuscles. Not only were the appearances entirely dif- 
ferent and the cartilage healthy—as otherwise ascertainable 
as well as from the known condition of the man and of the 
cause of his death—but the true nature of the lumps was 
made perfectly clear by examination with higher powers. 
When magnified to the extent of 600 diameters, the same 
relative appearance was preserved. The lumps in the basis- 
substance still varied in size from the limit of the visible to 
the magnitude of ordinary cartilage corpuscles ; but in all 
the larger lumps, differentiations were visible which ap- 
proached them in structure as well as in size to cartilage 
corpuscles. In some, one or more vacuoles, in others, a 
small or large nucleus, or even two nuclei, could be made 
out ; and a few (z. e.y occasionally one in some fields) 
showed irregular twin, or even triplet formation. 
The highest power threw a wonderful light upon these 
lumps. They were seen to be masses of living matter. The 
larger showed a network in their interior, some without and 
some with a nucleus, and the latter, when present, was 
sometimes homogeneous and sometimes reticulated. All 
the lumps, except the smaller, were surrounded by a dis- 
tinct light seam, through which radiating conical offshoots 
passed to the network in the basis-substance ; and all of 
them, even the smallest, sent delicate offshoots connecting 
them with that network, or were themselves part and parcel 
(z. e.t thickened points of intersection of the threads) of that 
network. 18 
Louis Elsbcrg. 
After having studied such a specimen, it was easy to in- 
terpret correctly the intrareticular granules seen in the alco- 
hol specimen represented in fig. 4. 
Fig. 5 shows a horizontal section of the thyroid cartilage 
of a robust, rather stout man, 48 years old, amplified 600 
diam. C C} cartilage corpuscles; G, granules or lumps 
scattered through the hyaline basis-substance ; F, fibrous 
portion of cartilage. 
Fig. 6 shows a field of the same specimen with an amplifi- 
cation of 1200 diam. C, cartilage corpuscle with distinct 
reticular structure surrounded by a dense basis-substance, 
the so-called capsule ; B, hyaline basis-substance with faint 
reticular structure; G, granules or lumps scattered through 
the hyaline basis-substance, connected with its reticulum. 
Section III.—Deductive* 
I.—Regarding the essential structure. 
Instead of being a mass of basis-substance in which a 
number of cartilage corpuscles are imbedded, hyaline car- 
tilage is a filigree of living matter, in the meshes of which 
lumps of basis-substance are imbedded. 
The difference between the former and the present view 
of the essential structure is a radical one. Of its full im- 
port I shall say more before the end of this Section. 
The number of investigators who have hitherto suc- 
ceeded in observing cartilage corpuscles sending offshoots 
into the basis-substance, is very small ; the number of those 
who have succeeded in tracing such offshoots far enough 
into the basis-substance to observe their forming a network 
is smaller still. As a rule, this network can be seen only 
very faintly in fresh specimens, and in specimens obtained 
either after hardening in chromic acid solution or by other 
methods ; but the staining with gold chloride, as practised 
by Heitzmann in 1872 brought it more plainly into view, as 
did also the alcohol method of examination devised by 
Spina in 1879. The fact that in the intervening seven 
years many observers have endeavored to see it and failed, 
Read in part before the New York Academy of Medicine, Nov. 17, 1881, and 
the New York Academy of Sciences, Nov. 21, 1881. 
* Reprinted from the Archives of Laryngolgy, Vol. iii, No. 1, Jan., 1882. Histology of the Cartilages of the Larynx. 
19 
may be due, as claimed by Spina, to the method employed. 
I certainly accord great merit to Spina for his discovery of 
the effect of the alcohol treatment. Not only is it far 
simpler in every way than the gold-chloride treatment, but 
being very easy of employment it is also invariably, so far 
as my experience hitherto has*gone, attended with suc- 
cess. Although many have failed with former methods, I 
venture to say that no one who follows the directions given 
by Spina, but will see the offshoots and the network. A dif- 
ferent question, however, than the mere seeing, is the correct 
interpretation. Spina himself does not follow out the logi- 
cal consequences of his observations, and Flesch, the au- 
thor of the latest publication on the subject, goes only so 
far as to say “ the offshoots are lost in the basis-substance, 
their continuation forms the cement-substance.” 1 Perhaps 
it needed the occurrence of so convincing a case as that re- 
corded in the present paper to place the matter of inter- 
pretation beyond dispute. At all events, so far as I am 
aware, no such clear demonstration of the correct interpre- 
tation of the structure of cartilage was ever had before. 
It is the case which figs. 5 and 6 illustrate. The basis- 
substance is crowded with lumps visible with even low pow- 
ers of the microscope without any other reagent than that 
the larynx had been preserved in chromic acid solution. 
The lumps are proved to be living matter by all the tests 
applied to them. Their identity in this respect and unin- 
terrupted filamentous and granular connection with the 
cartilage corpuscles on the one hand and with the granules 
and threads of the intercorpuscular network on the other, 
render doubt as to the interpretation impossible. In other 
words, we have in hyaline cartilage before us a mass of liv- 
ing matter arranged principally in the form of a granular 
network with large and small meshes; the large meshes con- 
tain chondrogenous firm basis-substance, the small meshes 
contain liquid, and with their usual but perhaps not essen- 
tial limiting layer constitute the cartilage corpuscles; 
thickened points of intersection of the filaments of the 
1 Loc cit.y p. 85. Though published in 1880, Flesch tells us that his work 
was finished by the end of 1878. Spina’s communication is not referred to. 
Flesch employed gold chloride and other methods. Louis Elsberg. 
20 
network constitute granules; comparatively smaller and 
larger lumps of living matter exist i) in the corpuscle as 
nucleoli and nuclei, the latter also usually already reticu- 
lated, and 2) in the basis-substance as granules and small 
corpuscles. 
II.—Regarding the so-called “juice routes” or “lymph- 
channels.” 
In my historical sketch I have given a very slight idea 
only of the many and elaborate researches to explain how 
nutrient juice might reach the cartilage corpuscle, isolated 
and enclosed in the hard basis-substance. But the truth 
being that the corpuscle is not isolated and so enclosed, 
much of the laborious investigation and ingenious reason- 
ing was inapplicable to the case. Of course I do not agree 
with those who think that cartilage contains a system of 
tubular or other-shaped channels, either with or without a 
lining membrane, or that it contains fibrils with interfibril- 
lar cement-substance; i. e., I do not agree with the advocates 
of either of these views in the sense in which they hold 
them. But I can see how, in the absence of the guiding 
knowledge which was supplied by Heitzmann’s great dis- 
covery of the arrangement of living matter, these differ- 
ent investigators arrived at their conclusions. The minute 
blocks of basis-substance are separated from each other, 
and at the same time joined together, by the living matter 
between them. This is the ground for the interpretation, 
on the one hand, that there exists a “ viscous-soft ” cement- 
substance in the intercorpuscular space capable of conveying 
juices, and, on the other, that there are canals which hold 
offshoots of the corpuscles. There is no good reason that 
I know of, for denying that, in addition to the living mat- 
ter, a certain amount of liquid may be circulating in such 
canals, i. e., in the interstices between blocks of basis-sub- 
stance in the intercorpuscular spaces (and perhaps also 
around the corpuscles themselves), and that such liquid may 
carry particles of foreign bodies as charcoal, carmine, cinna- 
bar, etc., far into the cartilaginous tissue ; but the liquid is 
not the essential portion of the contents of these canals, and 
if experimenters have succeeded in injecting them, the high Histology of the Cartilages of the Larynx. 
21 
pressure exerted upon the cartilage must have pushed 
aside or compressed the living matter. What the “juice 
routes ” are, is clear enough. When Spina had demon- 
strated that coloring matter is absorbed into cartilaginous 
tissue by means of the offshoots of the cartilage corpuscles, 
he need not have added the remark that proof is wanting 
that the coloring matter can pass through cartilage outside 
of such offshoots. Such proof will ever remain wanting. 
Every cartilage corpuscle, no matter how far removed it 
be from the source of nutrient supply, is more or less di- 
rectly connected therewith by means of the filigree of liv- 
ing matter; and this filigree itself constitutes the system of 
juice routes. 
III.—Regarding the intermixture of fibrous basis-sub- 
stance. 
The laryngeal, as well as the tracheal and the costal car- 
tilages, invariably contain, irregularly intermixed with hya- 
line basis-substance, a number of small or large portions of 
striated or fibrous basis-substance. Now, notwithstanding 
the constancy of this occurrence, all authors on the subject 
describe it as the result either of a degenerative process 
(Rheiner, Luschka, Schottelius, etc.), or of atrophy (Roki- 
tanski, etc.). According to Schottelius,1 the process com- 
mences in about the tenth or twelfth year of life, and im- 
mediately introduces profound destructions of the tissue. 
He says that the fibres are cylindrical filaments, arranged in 
general in a parallel direction, but somewhat interlaced. It 
seems to him that the formation of these fibres occurs when, 
in a degenerated basis-substance, the cartilage corpuscles 
perish ; perhaps by a coagulation process similar to that by 
which fibrine is formed. I shall show in the next chapter 
that Schottelius is mistaken in believing that the intermix- 
ture of striated basis-substance is not found until the 
approach of puberty. But even were he right, I should 
still contend that I have met with fibrous basis-substance in 
healthy as well as in diseased hyaline cartilage, and that 
investigation has not led me to believe that its presence 
1 Die Kehlkopfknorpel. Untersuchungen liber deren physiologische und 
pathologische Texturveranderungen. Wiesbaden, J. F. Bergmann, 1879, pp. 
20, 21. 22 
Louis Elsberg. 
necessarily results from a degenerative process. When, 
f. i., I find it in every healthy thyroid cartilage that I 
examine, I cannot but regard its development as normal. Is 
it not supererogation to look upon a phenomenon as degen- 
eration which is present—as is this, at least so far as my 
experience goes—in every individual, without exception ? 
I think the presence of fibrous portions must be considered 
normal in cartilages that appear healthy in every other 
respect; but I do not mean to deny the possibility of 
fibrous degeneration with other pathological conditions. 
In such a case of transformation there must be an inter- 
mediate state. All recent researches tend more and more 
to show that one tissue is never directly changed into an- 
other, and I cannot easily believe that what is first a 
hyaline basis-substance can directly become a fibrous one. 
Attempts to explain any such direct transformation have 
proved failures, I believe; and even with our present knowl- 
edge of the existence of living matter within the basis-sub- 
stance, it would be a difficult task to realize that the firm, 
solid, chondrogenous basis-substance can assume the stri- 
ated form. The least that would have to be admitted is a 
melting, i. e., a solution or softening of the basis-substance in 
certain territories of hyaline cartilage, with accompanying 
fusion of the living matter, and subsequent splitting up of 
the mass or masses of the latter into delicate spindles. 
That striated portions of hyaline cartilage have in the 
course of normal development arisen from medullary tissue, 
present in the foetal condition, seems far more probable. If 
this view be addpted, we need not assume with Schottelius 
and others, that all the blood-vessels found after puberty 
are necessarily newly formed, but that some have been pre- 
served from the infantile state. Indeed, I have met with 
appearances in my specimens which seemed strongly to in- 
dicate that some of the blood-vessels formerly present in 
the medullary tissue are now present between the fibres, 
while others have become obliterated, solidified, and trans- 
formed into the basis-substance themselves. 
IV.—Regarding the cell-doctrine and the bioplasson- 
doctrine. Histology of the Cartilages of the Larynx. 
23 
Just as the study of cartilage led to the cell-doctrine, 
which at the time of its establishment was a great advance 
in biological science, so the further study of cartilage sup- 
plied the basis for a generalization which is a further devel- 
opment, and must take the place, of the cell-doctrine. 
This is Heitzmann’s doctrine of living matter, or, as I have 
named it, the bioplasson-doctrine.* 
When the term “ cell ” was introduced, by Schleiden and 
Schwann, into histology, it was believed that the body on ul- 
timate morphological analysis was found to consist of a num- 
ber of minute vesicles or sacs, enclosing liquid contents in 
which is suspended a more solid body, the nucleus (this latter 
frequently containing a smaller similar body, the nucleolus, 
and this sometimes a still smaller one, the nucleolinus). For 
fully twenty years this idea has been known to be erroneous. 
Indeed, Goodsir nearly forty years ago—only a few years 
after Schwann had established the cell-doctrine and attrib- 
uted the so-called metabolic or vital power to the cell-mem- 
brane—had experimentally determined that the seat of the 
vital process of secretion is not in the vesicle as such, 
but in the so-called cell-contents ; Naegeli, in 1845, and Al- 
exander Braun, in 1851, had also shown the cell-wall to be 
comparatively unimportant ; and in 1857 Leydig had de- 
clared the “cell” to consist only of a soft substance en- 
closing a nucleus. Certainly, twenty years ago it was proved 
beyond dispute by Max Schultze, Beale, Hseckel, and others, 
that what was called a “ cell ” was not a vesicle, but es- 
sentially a jelly-like lump of living matter characterized by 
the presence of a nucleus; soon after, Robin, Briicke, 
Kiihne, Strieker, and others, conclusively showed that not 
even a nucleus is an essential constituent of an elementary 
organism; and biologists were compelled to transfer the power 
of manifesting vital properties to “living matter” instead 
of restricting this power to any definite form or element. As 
long ago as in 1861, Briicke proposed to discontinue the use 
of the word “cell” as being a misnomer and misleading, 
and offered as a substitute the expression “ elementary or- 
1 “ Notice of the Bioplasson-Doctrine.” Transactions of the American Medi- 
cal Association, vol. xxvi, 1875. 24 
Louis Elsberg. 
ganism.” Beale proposed, instead, the term “bioplast” to 
designate any definite mass of living matter, and Hseckel 
the term “plastid.”1 From the latter I devised the word 
“ plastidule ” as synonymous with ultimate molecule of the 
substance of living matter.2 Elementary living matter is 
called with Dujardin “ sarcode,” or with Von Mohl “pro- 
toplasm,” or with Beale “ bioplasm,” or, still better (be- 
cause it is a designation etymologically more nearly mean- 
ing living, forming matter), “ bioplasson.” 3 Of these four 
synonymous terms, protoplasm is best known, but has been 
used in other senses than to designate merely elementary 
living matter. I, therefore, think that bioplasson is to be 
preferred. 
According to Drysdale,4 Dr. John Fletcher of Edinburgh 
was the first6 who clearly abandoned the idea that the ma- 
terial elements of an organism require the addition “ of an 
immaterial or spiritual essence, substance, or power, general 
or local, whose presence is the efficient cause of life,” and 
who arrived at the conclusion that “it is only in virtue 
of a specially living matter, universally diffused and inti- 
mately interwoven with its texture, that any tissue or part 
possesses vitality.” He denied vitality to any gaseous or 
purely liquid fluid, and any hard or rigid solid ; and thought 
the only truly living matter consisted “of the gray matter 
of the ganglionic nerves, which he held to be universally 
1 The term “ Plastid ” was intended by Ilseckel to include every form-ele- 
ment, whether a cell in the older sense or a lump of living matter without an in- 
vesting membrane, and whether nucleated or unnucleated. For the nucleated 
plastid he retained the word cell or cyte, the unnucleated he called cell-like or 
cytode ; and, according to his precise discrimination, there are therefore four 
kinds of plastids, viz.: I. Gymnocytodes, or nude cytodes, i.e., unnucleated 
lumps of living matter without membrane ; 2. Lepocytodes, or covered cytodes, 
i. e., unnucleated plastids with investing membrane ; 3. Gymnocytes, or nude 
cells, i. e., nucleated plastids without investing membrane ; 4. Lepocytes, or 
covered cells, i. e., cells with a cell-wall. The progress of histology has made 
these distinctions for that department of less value than they were when origi- 
nally made, but they are still very useful for natural history purposes. 
2 See London Monthly Microscopical Journal, 1872, p. 182 ; “ Proceedings 
American Association for the Advancement of Science,” 1874 ; and “ On the 
Plastidule Hypothesis,” Proceedings of the same, 1876. 
3Of course, deadbioplasson is a contradiction in terms: bioplasson deprived 
of vitality is no longer bioplasson at all, but merely the chemical remains of 
what once was bioplasson. If this be remembered there will be no confusion, 
even if the word be used in describing tissues, etc., after death. 
4 The Protoplasmic Theory of Life, London, 1874. 
4 Rudiments of Physiology, Edinburgh, 1835. Histology of the Cartilages of the Larynx. 
25 
diffused, and the gray matter of the brain and spinal mar- 
row.” He described it as a “ nitrogenous, pulpy, translu- 
cent, homogeneous matter, yielding, after death, fibrin.” 
“ Chemical analysis, accordingly, must be considered as use- 
ful in showing us, not what such matter was composed of 
while it possessed vitality, but what it is composed of after- 
ward.” “ Not only is every vital action traced to molecular 
change, and to consumption and regeneration of this struc- 
tureless, semi-fluid matter, combined in a way entirely sui 
generis, but the initiation of these changes is brought by 
Fletcher into absolute dependence on stimuli, and all spon- 
taneity or autonomy is denied to matter in the living just as 
in the dead state.” 
As Fletcher’s work was published in 1835, several years 
before even the establishment of the cell-doctrine, we 
cannot but agree so far with Drysdale as to say that 
Fletcher has framed “ a hypothesis of the anatomical nature 
of the living matter which anticipates in a remarkable man- 
ner” its discovery ! In 1850, Cohn1 recognized the proto- 
plasm “as the contractile element, and as what gives to the 
zoospore the faculty of altering its figure without any cor- 
responding change in volume.” He concludes that proto- 
plasm “ must be regarded as the prime seat of almost all 
vital activity, but especially of all the motile phenomena in 
the interior of the cell.” In 1853 Huxley8 said “vitality, 
the faculty, that is, of exhibiting definite cycles of change 
in form and composition, is a property inherent in certain 
kinds of matter.” In 1855 Unger3 thought that “ the proxi- 
mate cause of the movements of the sap in the cells is to be 
sought neither in diosmosis, nor in the action of the nuclear 
vesicle, nor in any mechanical contrivance, such as cilia, but 
it lies rather in the constitution of the self-moving proto- 
plasm, which, as an especially nitrogenous body of the na- 
ture of that simple contractile animal substance called sar- 
code, produces the rhythmically advancing contraction and 
expansion.” 
I“Nachtrage zur Naturgeschichte des Protococcus pluvialis.” Nova acta 
Acad. Leop.-Carol., vol. xxii, part i., p. 605. 
a “ Review of the Cell-theory.” British and Foreign Medico-chirurg. Review, 
Oct., 1853. 
‘Anatomie und Physiologie der Pflanzen, 1855, pp. 280, 282. 26 
Louis Elsberg. 
In 1856 Lord S. G. Osborne discovered carmine staining, 
and distinguished by means of coloring it the living forma- 
tive matter from the formed material, a means which has 
borne important fruits in the discovery of Cohnheim’s stain- 
ing of living matter by gold chloride, and in that of Reck- 
linghausen’s staining all except living matter by silver ni- 
trate. 
In 1858, and in a number of later articles,1 Max Schultze, 
by showing that, as had been hypothetically supposed by 
Unger, the movements of the pseudopodia and the gran- 
ules are really produced by active contractile movements of 
the protoplasm, and by other observations, contributed 
much to the establishment of the theory of living matter. 
Haeckel has also for many years, and in various publica- 
tions,3 labored to maintain and extend the same theory, of 
which he thus expresses himself.3 “ The protoplasm or 
sarcode theory, that is * * * that this albuminous ma- 
terial is the original active substratum of all vital phenom- 
ena may, perhaps, be considered one of the greatest achieve- 
ments of modern biology, and one of the richest in results.” 
And says Drysdale4: “ If the grand theory of the one 
true living matter was, as we have seen, hypothetically ad- 
vanced by Fletcher, yet the merit of the discovery of the 
actual anatomical representation of it belongs to Beale, in 
accordance with the usual and right award of the title of 
discoverer to him alone who demonstrates truths by proof 
and fact. * * * The cardinal point in the theory of 
Dr. Beale is not the destruction of the completeness of the 
cell of Schwann as the elementary unit, for that was already 
accomplished by others. * * * But that, from the 
earliest visible speck of germ, up to the last moment of 
1 “ Ueber innere Bewegungs-Erscheinungen bei Diatomeen,” Muller s Ar- 
chiv, 1858, p. 330 ; “ Ueber Cornuspira,” Archiv f. Naturgesch., i860, p. 287; 
“ Ueber Muskelkorperchen und das was man eine Zelle zu nennen habe,” 
Reichert und Du Bois-Reymond's Archiv, 1861, p. I ; Das Protoplasma der 
Rhizopoden und der Pflanzenzellen, Leipzig, 1863. 
2 Monographic der Radiolarien, 1862, pp. 89, 116 ; “ Ueber den Sarcode- 
korper der Rhizopoden,” Zeitsch. f. IVissensch. Zodlogie, 1865, p. 342 ; Gen- 
erelle Morphologie, vol. i, pp. 269, 289. 
3 Monographic der Moneren,” Jenaische Zeitschft. f. Medicin und Natur- 
wissenschaft, 1868, iv, 1 ; translation in Quarterly Journal of Microscopical 
Science, London, 1869, vol. ix, p. 223. 
4 Loc. cit., p. 42, et seq. Histology of the Cartilages of the Larynx. 
27 
life, in every living thing, plant, animal, and protist, the at- 
tribute of life is restricted to one anatomical element alone, 
and this homogeneous and structureless; while all the rest 
of the infinite variety of structure and composition, solid 
and fluid, which make up living beings, is merely passive 
and lifeless formed material. This distinction into only two 
radically different kinds of matter, viz., the living or ger- 
minal matter and the formed material, gives the clue where- 
by he clears up the confusion into which the cell-doctrine 
had fallen, and gives the point of departure for the theory 
of innate independent life of each part, which the cell-theory 
had aimed at but failed to make good. The one true and 
only living matter—called by Beale germinal matter, or bio- 
plasm— is described as ‘ always transparent and colorless, 
and, as far as can be ascertained by examination with the 
highest powers, perfectly structureless ; and it exhibits those 
same characters at every period of its existence.’ * * * 
The living matter of Beale corresponds to the following his- 
tological elements of other authors : The viscid nitrogenous 
substance within the primordial utricle, called by Von Mohl 
protoplasm; the primordial utricle itself, in Naegeli’s sense 
of that term, viz., the layer of protoplasm next the cell-wall; 
the transparent, semi-fluid matter occupying the spaces and 
intervals between the threads and walls of those spaces 
formed by the so-called vacuolation of protoplasmic masses; 
the greater part of the sarcode of the monera, rhizopoda, 
and other low organisms ; the white blood-corpuscles, pus- 
corpuscles, and other naked wandering masses of living 
matter; the so-called nucleus of the secreting cells, and 
of the tissues of the higher animals, and many plant-cells ; 
the nuclei of the cells of the gray matter of the brain, spinal 
marrow, and ganglions, and the nuclei of nerve-fibres. The 
term of true living or germinal matter can never be given 
to the following parts, although to some of them the word 
protoplasm has been erroneously applied, viz., the cell-wall 
of plants or animals, however delicate or gelatinous; 
the threads or filaments and walls of the vacuoles within 
protoplasmic masses or cells; the wall of the primordial 
utricle ; the true fibrous, connective, elastic, bony, or other 28 
Louis Els berg. 
tissues generally included among the living parts of ani- 
mals ; even the proper contractile fibre of the muscles, the 
radiating fibres of the caudate nerve-cells, and the outer 
coat of those cells, besides the nerve-fibres in general; the 
hard parts of epithelial cells, and all liquid secretions ; the 
cilia ; the tissue of cuticle, hair, nails, horn, and all analo- 
gous parts in plants; the granules in sarcode; all coloring 
matter; and, lastly, all pabulum, including the fluid part of 
blood, lymph, and chyle, and corresponding matters in 
plants. In short, the name of bioplasm, given by Beale, or 
protoplasm (in a restricted sense as it will probably be ulti- 
mately accepted by biologists), as indicating the ideal living 
matter, cannot be given to any substance displaying rigidity 
in any degree, from the softest gelatinous membrane up to 
the hardest teeth-enamel ; nor to any thing exhibiting a 
trace of structure to the finest microscope ; nor to any liquid ; 
nor to any substance capable of true solution. Thus, ‘ noth- 
ing that lives is alive in every part,’ but as long as any indi- 
vidual part or tissue is properly called living it is only so in 
virtue of particles of the above-described protoplasm freely 
distributed among, or interwoven with the textures so close- 
ly that there is scarcely any part of an inch in size but 
contains its portion of protoplasm. Thus we see realized 
the hypothesis of Fletcher, that all * living action is per- 
formed solely by virtue of portions of irritable or living 
matter interwoven with the otherwise dead textures. Ac- 
cording to Beale, ‘of the matter which constitutes the 
bodies of man and animals in the fully-formed condition, 
probably more than four-fifths are in the formed and non- 
living state. All this was, however, living at an earlier period 
of existence.’ This is on an average, for some tissues con- 
tain much less living matter; the bones, for example, only 
and some textures, when old, not more than y^th.” 
I have made this long quotation from Drysdale’s book, 
because I am anxious to do full justice to Beale, and I 
could not find a statement of his views so succinct for quo- 
tation in his own writing. The objection, however, urged 
by Bastian to Beale is so very pertinent, that it must also 
find a place here, but I shall not dwell upon other points on Histology of the Cartilages of the Larynx. 
29 
which Beale differs from the bioplasson doctrine ; such as, 
that living matter exhibits the same characters at every 
period of its existence, and, that it is always perfectly 
structureless. “ It has always appeared to me,” says Bas- 
tian,1 “to be a very fundamental objection to his theory, 
that so many of the most characteristically vital phenomena 
of the higher animals should take place through the agency 
of tissues—muscle and nerve, for instance—by far the 
greater part of the bulk of which would, in accordance with 
Dr. Beale’s view, have to be considered as dead and inert.” 
In 1873 the morphological knowledge of living matter 
became exact. In that year Heitzmann discovered the 
manner in which bioplasson is arranged throughout the 
body, and announced the fact that what had until then 
been regarded as separate form-elements in a tissue are 
really interconnected portions of living matter; that not 
only are there contained no isolated unit-masses in any one 
tissue, but no tissue in the whole body is isolated from the 
other tissues ; and that the only unconnected particles of 
living matter are the corpuscular elements of liquids, such as 
blood, sperm, saliva, pus, etc., and so-called wandering cor- 
puscles; so that, to use his own words : “The animal body 
as a whole is a connected mass of protoplasma in which, in 
some part, are imbedded isolated protoplasma-corpuscles 
(wandering corpuscles, colorless and red blood-corpuscles) 
and various not-living substances (glue-giving and mucin- 
containing substances in the widest sense, also fat, pigment- 
granules, etc.).” This announcement marked the commence- 
ment of a new era in biology. 
Heitzmann had “ investigated the condition of living mat- 
ter in so-called infusions.” There had at first appeared “a 
number of very minute granules, just perceptible to the 
highest powers of the microscope,” granules that were 
homogeneous, yellowish, shining, and motionless. Such a 
granule represents the juvenile phase of living matter; 
growing larger in size it becomes vacuolized, i. e., shows “ a 
central hole, enclosed on all sides by a yellowish shining 
1 The Beginnings of Life: being some account of the nature, modes of origin, 
and transformations of lower organisms. London, 1872, vol. i., p. 155. 30 
Louis Elsberg. 
substance.” “ The cavities in the interior of the shining 
lumps, apparently filled with some fluid, differ in their 
refracting power from the surrounding mass ; they show a 
slight rosy color.” Still later, “ some granules look as if 
perforated by vacuoles, like a sieve ; the differentiation 
between the two substances within the lump—the yellowish 
shining and the colorless rosy-refracting—has apparently 
advanced.” Vacuolation produces the appearance at first 
of a trabecular framework, and then of a filamentous net- 
work in the interior of the lump. “ Such a minute trans- 
parent corpuscle, floating in the water of the infusion, con- 
stantly changes its outlines by throwing out offshoots or 
processes, mostly in the form of hyaline flaps ; and we are 
sure we have an amoeba before us.” Heitzmann discovered 
that the living matter as seen in an amoeba is not without 
structure, as had, before his accurate investigations, been 
supposed ; and that its structure, in all cases when devel- 
oped, is that of a network, in the meshes of which the 
bioplasson fluid, or the not-contractile, not-living portion of 
the organism, exists. When there is a nucleus, which may 
be either homogeneous (/. e., compact) or vacuolized or 
reticulated, it is connected by delicate threads with the 
extranuclear network; nucleoli and nucleolini inside of 
the nucleus, as well as granules outside, are portions of 
living matter : sometimes in lump, sometimes mere points 
of intersection of the threads constituting the intranuclear 
and extranuclear living networks, sometimes terminals of 
section of such threads, as first explained by Eimer,1 and 
after him by Klein.2 
Heitzmann discovered that what is true of the structure of 
bioplasson in the amoeba, where a single small unit-mass of 
living matter constitutes the entire individual, is true also 
1 “ Weitere Nachrichten liber den Bau des Zellkerns.” Archiv f. mikrosk. 
Anatomie, xiv, 1877, p. 103. 
3 “ Observations on the Structure of Cells and Nuclei,” Quarterly Journal 
of Alicroscopical Science, Jan., 1879, p. 128. “The intranuclear as well as the 
intracellular network having, of course, three dimensions, includes fibrils that 
lie in the two dimensions of the plane of the field of the microscope, as well as 
fibrils placed vertically to it. The former appear, of course, as fibrils : but, I 
should like to ask, as what do the latter appear, i. e., those situated vertically. 
Clearly as dots, because they are seen endwise; and for obvious reasons most of 
them lie in the nodes of the network.” Histology of the Cartilages of the Larynx. 
31 
of the structure of bioplasson of all, even the highest, living 
organisms. 
It was known that the ideas originally connected with 
the term “ cell ” were erroneous, and that the essential con- 
stituent of “cells” was living matter individualized into 
small distinct masses; the existence in such form-elements 
of granules, aside from membrane, nucleus, nucleolus, and 
nucleolinus, was known ; “ thorns” and offshoots, processes 
and prolongations from them had even been occasionally 
observed ; and Frommann1 is even said5 to have observed 
“ clear shining fibrils proceeding from the nucleolus joining 
themselves to others proceeding from the nucleus and pro- 
toplasm,” and to have believed “ that the granules of the 
nucleus and protoplasm are the nodal points of a very fine 
fibrous network, from which fibrils go off and leave the cell.” 
But all this knowledge was fragmentary until Heitzmann 
demonstrated not only that membrane, nucleus, nucleolus, 
granules, and threads are really the living contractile matter, 
but also 1) that this matter is arranged in a net work, contain- 
ing in its meshes the non-contractile matter, which is trans- 
formed into the various kinds of basis-substance, character- 
izing different tissues; and 2) that the tissue-masses of bio- 
plasson throughout the whole body are interconnected by 
means of fine threads of the same living matter. 
Unless these facts of Heitzmann’s discovery are accepted, 
there cannot be urged much against the continued use of the 
word “cell,” misnomer though it be. Ranke,3 after speaking 
of the “ cell-wall,” “cell-nucleus,” etc.,says: “Of these com- 
ponent parts of the cell, one or other may be wanting with- 
out the totality ceasing to be a cell. . The nucleoli, the cell- 
wall, or the nucleus may be wanting, and yet we must desig- 
nate the microscopic form a cell, or elementary organism.” 
Now, although Drysdale thus comments upon this quota- 
tion, viz.: “ If any one choose to describe a gun-barrel as a 
stockless gun without a lock, he is free to do so ; but what 
1 “ Untersuchungen ttber die normale und pathologische Anatomie des Riick- 
enmarks,” 2 Theil, Jena, 1867. 
2 As I have at present no access to Frommann’s pamphlet, I must content 
myself with this expression and make the following citations from Tyson, The 
Cell-Doctrine : its history and present state. Philadelphia, 1878, p. 117. 
3 “ Physiologie, 1872,” quoted by Drysdale, loc. cit., p. 104. 32 
Louis Els berg. 
good purpose can it serve ? Or is there even any fun in it ? 
The truth is, this clinging to the mere name of the cell- 
theory by the Germans seems to arise from a kind of per- 
verted idea of patriotism and of pietas toward Schwann 
and Schleiden.” Nevertheless, I think Tyson1 has the 
better of the argument, in saying: “ The word has, how- 
ever, become so intimately associated with histology, that 
it is doubtful whether it will ever fall into disuse, nor does 
it much matter, so long as correct notions of the elemen- 
tary part are obtained.” If there were any separate and 
distinct “elementary part,” it certainly would matter little or 
nothing whether it were called “ cell ” or by any other name, 
provided the name be properly defined and agreed upon. 
It is not against the name but against the idea of any 
isolated individualized form-element that the objection lies. 
Virchow maintains* “that the cell is really the ultimate 
morphological unit in which there is any manifestation of 
life, and that we must not transfer the seat of real action to 
any point beyond the cell.” Against this statement nearly 
every author nowadays protests, and insists that vital 
power must be transferred from the “ cell ” to “ living mat- 
ter ” ; yet, after all, the disagreement, though ever so 
strenuously declared, is a mere verbal one: so long as both 
parties hold that “ every higher animal presents itself as a 
sum of vital unities” — no matter what these unities are 
called or how defined. Haeckel, one of the most avowed 
advocates of “ the protoplasm or sarcode theory,” clings to 
Virchow’s politico-physiological comparison, that every 
higher organism is like an organized social community or 
state, in which the individual citizens are represented by 
the “ cells ” [no matter how he may define these], each 
having a certain morphological and physiological autonomy, 
although on the other hand interdependent and subject to 
the laws of the whole. Heitzmann’s views necessitate the 
comparison of the body to a machine, such as a watch or a 
steam-engine, in which, though there are single parts, no 
1 Loc. cit. (see footnote 2, on p. 31), p. 128. 
2 Die Cellularpathologie in ihrer Begriindung auf physiologische und patho- 
logische Gewebelehre, Berlin, 1858, p. 3. (Translation by Chance, London, 
1859. P- 3-) Histology of the Cartilages of the Larynx. 
33 
part is at all autonomous, but all combine to make up one 
individual. According to Haeckel the body is composed of 
colonies of amoebae ; according to Heitzmann the body is 
one complex amoeba.1 
There is no better test of the truth of the bioplasson doc- 
trine than the structure of hyaline cartilage. If hyaline 
cartilage consists, as “is generally believed,” of “a homo- 
geneous ground-substance in which are closed cavities 
harboring nucleated cells,”2 the bioplasson doctrine is erro- 
neous. If it merely contains lymph- or juice-channels, no 
matter what their character, whether open or closed, 
whether lined or unlined, whether in “homogeneous basis- 
substance,” or “between layers of cells,” or “in cement- 
substance,” the bioplasson doctrine is erroneous. On the 
other hand, the more or less convincing observations by 
Frommann, Heitzmann, Hertwig, Thin, Prudden, Spina, 
and Flesch, that cartilage-corpuscles have offshoots, or 
prolongations, extending more or less far into the basis- 
substance, even that these prolongations anastomose with 
each other and connect neighboring corpuscles, are by no 
means sufficient to establish the doctrine. Of the three 
authors whose accounts are most satisfactory, Heitzmann 
himself, though influenced by these observations, did not 
depend upon them, but upon confirmatory investigations 
of other tissues ; Spina—though he might logically have 
followed the consequences of his observation and reason- 
ing—-is entirely silent on the subject; and Flesch distinctly 
repudiates the doctrine thus: “The views here presented 
agree in many respects with those of Heitzmann. It shall 
be reserved for an intended speedy continuation of these 
1 Since this was written, even Huxley, the popular champion of protoplasm as 
the physical basis of life, who, as Bastian (loc. cit., p. 58) says, so far back as 
in 1853, had put forth a powerful remonstrance against the then all-prevalent 
“ cellular theory ” of organization, has delivered an address before the Inter- 
national Medical Congress in London, August 9, 1881, in which he used the 
following language : “ In fact, the body is a machine of the nature of an army, 
not of that of a watch, or of a hydraulic apparatus. Of this army, each cell is 
a soldier, etc., etc., etc. 
2 This statement of the general belief is quoted from the introductory para- 
graph of Thin’s memoir “On the Structure of Hyaline Cartilage” (loc. cit.), 
in which Thin’s own views are laid down to the effect “ that layers of cells 
epithelial in arrangement exist in the substance of cartilage,” “ that both the 
stellate and the parallel systems of lymph-channels exist,” etc. 34 
Louis Elsberg. 
investigations to enter into a discussion of questions con- 
cerning the general cell-doctrine. Only this I must empha- 
size here in opposition to Heitzmann, and in accord with 
Max Schultze, that the independence (selbststiindigkeit) of 
the cell in the finished tissues does not suffer by the por- 
trayed arrangement, although its connection, both with the 
surrounding basis-substance and, through the latter, with 
neighboring cells, is more intimate than, perhaps, appears 
from customary descriptions.” 1 
The observations I have recorded in the second section, 
and especially those illustrated in figs. 4, 5, and 6, admit of 
but one interpretation, and that an interpretation favorable 
to the bioplasson doctrine. It is unnecessary to more than 
mention that although I have placed on record so few, I 
have made many different examinations, under many differ- 
ent circumstances, and with varying powers of amplification. 
I need occupy myself here with only the two fields drawn 
in figs. 5 and 6, with an amplification of 600 and 1200 
respectively. The remarkable specimens from which they 
are taken show more conclusively than it was ever before 
shown what the structure or constitution of hyaline carti- 
lage really is. I have explained this sufficiently under the 
proper heading. Its full significance now appears in its 
corroboration of the bioplasson doctrine. To be able to 
uphold the cell-doctrine, cartilage would have to be, using 
a homely comparison, like a cake composed of hard dough 
with raisins. No matter how widely we may extend the 
definition, to remain within the boundary of that doctrine 
this metaphor must be applicable. Innumerable pains- 
taking researches have led to various modifications of 
notions entertained regarding the structure of these two 
constituents and their relation to each other. As we have 
seen, especially by the most recent publication on the 
subject, viz., that of Flesch, the acceptation of the exist- 
ence in the dough of cleavage in certain directions, of 
interlaminary and interfibrillar spaces, and of offshoots, 
even ramifying prolongations of the raisin-substance, or, 
at all events, of an ingredient of the raisins, is held to be not 
1 Loc. cit.y p. 86. Histology of the Cartilages of the Larynx. 
35 
incompatible with that doctrine. If, however, we can repre- 
sent cartilage as a filigree or framework of raisin-substance, 
with here and there larger and smaller accumulations of 
this substance, in the meshes or interspaces of which frame- 
work blocks of dough are imbedded, certainly the funda- 
mental view of the ultimate construction of the tissue is 
changed, and we are no longer in accord with the cell- 
doctrine, even though we be inclined to use that term in 
the widest possible sense. 
In regard to a name as a substitute for the term “cell,” I 
would say that all corpuscular masses may be called, simply, 
corpuscles. For all the accumulations of living matter 
within the ordinary fields of basis-substance, but more espe- 
cially for those smaller masses which, having as yet devel- 
oped neither a network structure nor much vacuolation, 
are still homogeneous, or nearly so,—I am quite willing to 
adopt either the designation of “ plastids,” proposed by 
Haeckel, or that of “bioplasts,” proposed by Beale. Perhaps 
it would be best to restrict the word “ bioplast ” to a small 
mass of living matter exhibiting no differentiation, and to 
distinguish from it as “ plastid ” the larger mass showing 
an interior structure more or less like the fully developed 
corpuscle. Thus, I would always use the term “ plastid ” in 
the place of “ cell.” 
The result, then, of my investigations as to the structure 
of cartilage is that in this tissue, beyond the possibility of 
a doubt, the living matter is arranged in the form of a net- 
work containing in its meshes the non-contractile matter. 
How is it with regard to the other proposition of the bio- 
plasson doctrine, viz., that the living matter of the different 
tissues is interconnected? Examinations with high powers 
of such a specimen as that represented in fig. i, showing 
the perichondrium, of horizontal sections through the larynx, 
or the neck, with skin and more or less of other tissues in- 
cluded,—and here I desire to say a word in appreciation of 
Dr. Carl Seiler’s section-cutter, and of this gentleman’s 
kindness in preparing some very fine sections for me,— 36 
Louis E Is berg. 
enable me to answer this question to the effect that fine 
filaments of living matter pass from one tissue to another 
in connection with the network of living matter in each. 
The details of such examinations are reserved for another 
publication. ARCHIVES OF LARYNGOLOGY 
EDITED BY 
LOUIS M. D., 
N ' NEW YORK, 
IN CONJUNCTION WITH 
J. SOLIS COHEN, M. D., 
PHILADELPHIA, 
FREDERICK J. KNIGHT. M. D., 
BOSTON, 
GEORGE M. LEFFERTS, M. D., 
NEW YORK, 
pud Dr. J. Boeckel, Strassburg ; Prof. Burow, Kttnigsberg; Dr. Foulis, Glasgow; Prof. Ger- 
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