My examinations of the ova and embryos were chiefly made from the “sediment” sent in a special glass tube. The fully formed embryos were 1/125″ in length by 1/2500″ in breadth. They each showed a double skin, the outer envelope in the more advanced specimens leaving clear spaces at either end of the body, resulting from commencing ecdysis. I saw no trace of intestinal tube, but a central line of condensation marked an early differentiation of the somatic granular contents. The less advanced embryos were mostly enclosed in a chorional envelope, the smallest free embryos measuring only 1/200″ in length by 1/3000″ in breadth. These had no double contour. The ova, whose yolk-contents were still in various stages of cleavage, gave an average long diameter of 1/900 to 1/1000 of an inch.
Such are the facts I made out, and they enabled me to amend the characters of the species.
As regards nomenclature, I associated Dr Bancroft’s name with the sexually-mature worm as being in harmony with the binomial method and little calculated to mislead; moreover, it helped to fix both the source and date of the discovery (Brisbane, Dec. 21st, 1876). The use of this nomenclature detracts nothing from the high merits of Lewis, who first named the immature worm Filaria sanguinis hominis. As it now turns out, both Dr Salisbury and myself had previously been made acquainted with the young of Filaria Bancrofti; but it was reserved for Lewis to discover the hæmatozoal character of the embryos of this worm, and actually to take them from the blood. It was a singular circumstance, that when I was engaged in treating my little African patient for trematode hæmatozoa, it never once occurred to me that the numerous nematoid embryos mixed with the Bilharzia ova were hæmatozoal. As before remarked, it was alleged that my patient had passed worms two or three inches long by the urethra. I therefore concluded that these were the parents of the eggs and embryos, and that all of them were urinary. The inference was wrong, but it has instructively shown how near one may go towards a great discovery without really making it. As regards the larvæ, notwithstanding some slight differences in regard to size and so forth, I have little hesitation in saying that all the embryo forms severally described by Salisbury, by myself, by Lewis, Sonsino, Wucherer, Crévaux and Corre, Silva Lima, Bancroft, Manson, and others, are referable to one and the same species.
Into the clinical bearings of this subject it is impossible for me to enter at any length, but I may remark that these parasites appear to be associated with, if not actually the cause of, several distinct morbid conditions. To one of these Bancroft has given a separate name (Helminthoma elastica). This is a highly elastic form of growth to which I have already alluded under the title of “lymphatic abscess of the arm.” In the first valuable report on Hæmatozoa, by Dr Patrick Manson, of Amoy, China, this careful observer gives interesting particulars of no less than fifteen cases in which hæmatozoa were found. Two of these patients had Elephantiasis scroti, two had lymph-scrotum, two were lepers (one having scrotal disease), two had enlarged inguinal glands, one had anasarca; and of the remaining six, spoken of as having no concomitant disease, one had enlarged glands and abscesses, and another suffered from marked debility. It would thus appear that what is ordinarily termed “good health” is rarely associated with a hæmatozoal condition of the blood in the human subject. The cases given by Lewis and Manson, where absolutely no recognisable disease existed, must be regarded as exceptional. Disease, moreover, may exist without any palpable symptoms being exhibited by the “bearer,” and thus perhaps it was with the hæmatozoal dogs of Gruby and Delafond to which I shall again have occasion to allude. Even those animals that carried upwards of two hundred thousand microscopic Filariæ in their blood appeared to suffer no inconvenience whatever.
In the autumn of 1877 Dr Da Silva Lima published an article in the ‘Gazeta Medica da Bahia,’ in which he dwelt upon the labors and merits of Wucherer, and, judging from an omission in one of my memoirs, he supposed that I had insufficiently acknowledged Wucherer’s claims. A translation of this article appeared in the ‘Archives de Médicine Navale,’ with an important appendix by Dr le Roy de Méricourt. In this addendum the French savant showed that the omission on my part was unintentional, and had been corrected by me in a later memoir. Not only had I been amongst the earliest in England to enforce Wucherer’s claims in respect of the micro-Filariæ, but I had first announced his discoveries in connection with Anchylostoma duodenale. In my translation of Wucherer’s memoir (‘Ueber die Anchylostomum Krankheit’) I spoke of the melancholy satisfaction I had in knowing that the memoir in question was “among the last that appeared from the pen of that gifted and amiable physician.” Some notice of Dr Lima’s paper and its appendix by Dr A. le Roy de Méricourt appeared in the ‘Lancet’ for Jan. 5th, 1878, and I also published a full translation of it, with explanatory notes, in the ‘Veterinarian’ for Feb., 1878. Later on, in the ‘Lancet’ (March 23rd, 1878), Dr Da Silva Lima published an interesting letter correcting a misconception that had incidentally arisen in the mind of a commentator (on the Helminthological work of 1877), and at the same time he pointed to the original facts connected with the discovery of Wucherer’s Filaria. As my views are in perfect accord with those of Dr Da Silva Lima, I can only regret that errors of interpretation should have crept into the discussion. Dr Lima honorably recognises the nomenclature (Filaria Bancrofti) which I proposed for the adult worm, and only claims for Wucherer that which is fairly due.
On the 4th of January, 1878, I received from Dr Patrick Manson a manuscript in which he announced the discovery of the larvæ of Filaria sanguinis hominis in the stomach of mosquitoes. Already, in April, 1877, Dr Bancroft had informed me of his expectation of finding that these insects sucked up the larvæ of the Filaria whilst engaged in their attacks on man. Dr Bancroft’s supposition was a very natural one, but it remained for Manson to make the actual discovery of the existence of human hæmatozoa, or parasites that had been such, within the stomach of Culex mosquito. I lost no time in making the principal facts public (‘Lancet,’ Jan. 12th, 1878). Dr Manson at the same time forwarded for publication a record of thirty-five additional cases of hæmatozoa occurring in Chinese subjects, together with additional particulars of one of the cases already published in the ‘Customs Gazette.’ These were afterwards published as separate contributions in the ‘Medical Times and Gazette.’ Dr Manson likewise forwarded materials for a paper entitled “Further Observations on Filaria sanguinis hominis.” In this communication he gave an analysis of the cases (sixty-two in all) in which he had observed the hæmatozoa, and he added valuable statistical evidence as to the prevalence of Filariæ in the Amoy district, dwelling especially on the influence of age, sex, and occupation in determining the presence of the parasite. He also described the morbid states with which these entozoa were commonly associated.
On the 7th of March, 1878, I formally communicated to the Linnean Society a detailed account of Manson’s investigations relating to the metamorphoses undergone by the Filariæ within the body of the mosquito. In this paper Manson pointed out that the female mosquito, after gorging itself with human blood, repairs to stagnant water for the purpose of digesting the blood, and also for the purpose of depositing its eggs. During this period, which lasts four or five days, the Filariæ undergo remarkable changes. Subsequently, in a more perfect state, they escape into the water, and in this advanced stage they are conveyed to the human body along with the water as drink. Dr Manson persuaded a Chinese, whose blood was previously ascertained to abound with Filariæ, to sleep in a “mosquito house.” In the morning the gorged insects were captured and examined under the microscope. A drop of blood from the mosquito was found to contain 120 Filariæ, but a drop taken from the man’s hand yielded only some thirty specimens. Further stages of development are accomplished within the human host, ending in the sexual maturity of the parasite. After fecundation successive swarms of embryos are discharged by the female worm, a part of whose progeny eventually gains access to the blood.
Before I proceed to summarise the whole body of facts I must in the next place state that Manson and myself contributed a joint communication to the Medical Society of London on the 25th of March, 1878. In this memoir I especially dealt with the question of priority in connection with the discovery of the adult worm. I then restated that the adult parasite was discovered by Dr Bancroft on December 21st, 1876. The discovery was verified by Dr Lewis on August 7th, 1877, by Dr Silva Araujo October 16th, 1877, and by Dr F. dos Santos November 12th, 1877. I gave these dates unhesitatingly, without, however, in any way prejudicing the question already raised in respect of the identity of the worms found in each case. My own mind was fully made up on that point, and affirmatively so. Dr dos Santos’ find was made in conjunction with Dr J. de Moura in a case of lymphatic abscess of the arm. Clinically viewed, the case published by Dr Araujo must be regarded as unique. Not only were adult and embryonic Filariæ found in the same patient, but, what was far more surprising and interesting, the patient displayed in his own person several of the disorders hitherto found apart; and he was more than once attacked by one or two of the diseases. He experienced a first attack of chyluria three years ago, then attacks of craw-craw commencing a year ago, the latter being attributed to bathing in a particular lagoon. He had a second attack of chyluria six months back, at which time lymph-scrotum appeared, and also scrotal elephantiasis. Dr Bourel-Roncière pronounced this case to be unique, and attributed nearly all the disorders to the presence of Wucherer’s embryonic Filariæ. In a very elaborate analysis of and commentary on Dr da Silva Lima’s second memoir, Dr Bourel-Roncière warmly claims for Wucherer the supreme honor in all these discoveries. A number of affections hitherto regarded as distinct, and all of which appear to be due to the action of Filariæ, are regarded by Dr Bourel-Roncière as mere phases of one and the same disorder. This affection he terms Wucherer’s helminthiasis. Dr Manson had indeed arrived independently at a similar conclusion, and I am confident that Wucherer, were he alive, would in this particular aspect of the question be the last to claim priority either to Lewis, to Bancroft, or to Manson.
In this place I may observe that Dr Pedro S. de Magalhães, of Rio de Janeiro, detected free microscopic nematodes in the potable waters of Rio (agua da Carioca), which from their similarity he supposes may have some genetic relation with Filaria Bancrofti. In this opinion I cannot share.
As regards the metamorphoses of the embryo, Manson states that for a little while after gaining access to the stomach of the mosquito the embryo undergoes no change (Fig. [40], a). In a very few hours changes commence, resulting in wider separation of the outer skin and an appearance of transverse markings on the body within (b). In the next stage oral movements occur; the striation becomes more marked, and the outer envelope is cast off (c). Then the striated lines disappear and a dotted appearance is substituted (d). From this condition the embryo passes to what Manson calls the chrysalis stage, in which nearly all movement is suspended and the large spots gradually disappear (e, f, g, h, i, j, k). The tail continues to be flexed and extended at intervals and the oral motions cease. By the close of the third day the embryo becomes much shorter and broader; but the finely pointed tail retains its original dimensions, projecting abruptly from the sausage-shaped body (m, n). Large cells next appear in the interior of the body, and by a little pressure one may detect indications of a mouth (o, p, q, r). At this period the embryo begins to elongate, and at the same time to diminish in width; but the growth takes place chiefly at the oral end of the body. The mouth becomes four-lipped, open, and funnel-shaped, and from it a delicate line can be distinctly traced passing to an opening near the caudal extremity, the tail itself gradually disappearing (s, t). Speaking of the most advanced stage Manson says:—“A vessel of some sort is seen in the centre running nearly the whole length of the body and opening close to one extremity. This end is slightly tapered down and is crowned with three or perhaps four papillæ, but whether this is the head or tail, and whether the vessel opening near it is the alimentary canal or the vagina, I cannot say.” Now it is quite evident, I think, from Manson’s figures that he has here faithfully represented the head and tail, the former (u) to the left, the latter (v) to the right. In his manuscript (from which I am now quoting) there is no special reference to these two figures; but it is easy to see that these terminal sections of the body of the advanced embryo closely correspond with the head and tail of the adult worm (Filaria Bancrofti). The curved line passing to the left (u) evidently indicates the commencement of the partially-formed vagina.