Pearls & Parasites

The last few years of the nineteenth and the first few years of the present century are marked in the annals of medicine by a great increase in our knowledge of certain parasitic diseases, and, above all, in our knowledge of the agency by which the parasites causing the diseases are conveyed from host to host.

Chief among these agencies in carrying the disease-causing organisms from infected to uninfected animals are the insects, and, amongst the insects, above all the flies. Flies—e.g., the common house-fly (Musca domestica)—can carry about with them the bacillus of anthrax, and, if brought into contact with a wounded surface, may thus set up an outbreak of woolsorter’s disease. Flies, ants, and other even more objectionable insects, are not only capable of disseminating the plague bacillus from man to man, and from rat to man, but they themselves fall victims to the disease, and perish in great numbers. They are active agents in the spread of cholera, and the histories of the South African and Cuban wars definitely show that flies play a large part in carrying the bacilli of enteric fever from sources of infection to the food of man, thus spreading the disease. They are also accused of conveying the inflammatory matter of Egyptian ophthalmia, and of the ‘sore-eye,’ so common in Florida, from one human being to another.

The diseases already mentioned are caused by bacteria. But flies also play a part in the conveyance of a large number of organisms which are not bacteria, but which, nevertheless, cause disease, and cause it on the largest scale.

Of all the twenty-two orders into which the modern entomologist divides the class Insecta, that of the Diptera, or true flies, is, perhaps, the easiest to recognize, for it is characterized by one very obvious feature, the presence of the fore-wings only. The hind-wings are replaced by a pair of small-stalked, club-shaped ‘balancers,’ which are readily visible in some kinds of fly—e.g., the daddy-long-legs—but in others are by no means conspicuous. Thus it is an easy matter to determine whether an insect be a fly or not. To determine what particular kind of fly it be is, however, a very different affair. At present some forty thousand species of Diptera are known, and have been more or less completely described or figured; and Mr. D. Sharp estimates that this number is ‘only a tithe of what are still unknown to science.’ Further, the group has been rather neglected. Flies, speaking generally, are neither attractive in their appearance nor engaging in their habits, and it is a cause for no astonishment that entomologists have preferred to work at other groups.

In considering the part played by flies in disseminating diseases not caused by bacteria, we can neglect all but a very few families, those flies which suck blood having alone any interest in this connexion.

From the point of view of the physician, by far the most important of these families is the Culicidæ, with over three hundred described species and five sub-families, of which two, the Culicina and the Anophelina, interest us in relation to disease. The gnats or mosquitoes—the name is indifferently used, and has no scientific application—are amongst the most graceful and most beautiful insects that we know, but they have been judged by their works, and undoubtedly are unpopular, and we shall see that this unpopularity is well deserved. Gnats belong both to the genus Culex and to the genus Anopheles. The genus Culex, from which the order takes its name, includes not only our commonest gnat, often seen in swarms on summer evenings, but some hundred and thirty other species. Members of this genus convey from man to man the Filaria nocturna, one of the causes of the widely-spread disease filariasis, one variety of which is the elephantiasis, so common in parts of the tropics. In patients suffering from this disease minute embryonic round-worms swarm in the bloodvessels of the skin during the hours of darkness. Between six and seven in the evening they begin to appear in the superficial bloodvessels, and they increase in number till midnight, when they may occur in such numbers that five or six hundred may be counted in a single drop of blood. After midnight the swarms begin to lessen, and by breakfast-time, about eight or nine in the morning, except for a few strayed revellers, they have disappeared from the superficial circulation, and are hidden away in the larger bloodvessels and in the lungs.

In spite of their incredible number—some authorities place it at thirty to forty millions in one man—these minute larval organisms, shaped something like a needle pointed at each end, seem to cause little harm. It might be thought that they would traverse the walls of the bloodvessels and cause trouble in the surrounding tissues; but this is prevented by a curious device. It is well known that, like insects, round-worms from time to time cast their skins, and the young larvæ in the blood cast theirs, but do not escape from the inside of this winding-sheet; and thus, though they actively wriggle and coil and uncoil their bodies, their progress is as small and their struggles as little effective as are those of a man in a strait-waistcoat.

The causes of the periodicity of the appearance of these round-worms in the superficial bloodvessels are not completely understood, but they appear to have more relation with the usual sleeping hours of humanity than with day and night. In individuals who sleep by day and work by night the Filaria nocturna is found in the bloodvessels of the skin during the day. Thus, whilst between 5 p.m. and 7 or 8 a.m. the vessels of the skin of Cox the Hatter would be well peopled by the round-worms, they would only come to the surface in Box the Printer during the daytime, whilst he was sleeping in the lodgings of Mrs. Bouncer.

One reason of the normal appearance of the creatures in the blood at night is undoubtedly connected with the habits of its second host, the gnat or mosquito. Two species are accused of carrying the Filaria from man to man—Culex fatigans and Anopheles nigerrimus. Sucked up with the blood, the round-worms pass into the stomach of the insect. Here they appear to become violently excited, and rush from one end to the other of their enveloping sheath, until they succeed in breaking through it. When free, they pierce the walls of the stomach of the mosquito, and come to rest in the great thoracic muscles. Here the Filarias rest for some two or three weeks, growing considerably, and developing a mouth and alimentary canal; thence, when they are sufficiently developed, they make their way to the proboscis of the mosquito. Here they lie in couples, and it would be interesting to determine whether these couples are male and female. Exactly how they effect their exit from the mosquito and their entrance into man has not yet been accurately observed, but presumably it is during the process of biting. Only inside man they work their way to the lymphatics, and very soon the female begins to pour into the lymph a stream of young embryos, which reach the bloodvessels through the thoracic duct. It is, however, the adults which are the source of all the trouble. They are of considerable size, three or four inches in length, and their presence, by blocking the channels of the lymphatics, gives rise to a wide range of disease, of which elephantiasis is the most pronounced form. We can consider later how the disease can be averted by keeping down the number of gnats and by preventing their access to infected patients.

We now pass to the second of the diseases carried by gnats, that of malaria.

The parasite which causes malaria is a much more lowly organized animal than the Filaria. It is named Hæmamœba, and it, too, is conveyed by an insect, and, so far as we know, by one genus of mosquito only, the Anopheles. Hence, from the point of view of malaria, it is important to know whether a district is infected with Culex or Anopheles. The former is rather humpbacked, and keeps its body parallel with the surface it is biting, and its larva hangs at an angle below the surface of the water, by means of a respiratory tube. Anopheles, on the other hand, carries its body at a sharp angle with the surface upon which it rests, and its larva lies flat below the surface-film and parallel with it. The malarial parasite lives in the blood-cells of man, but at a certain period it breaks up into spores, which escape into the fluid of the blood, and it is at this moment that the sufferer feels the access of fever. The presence and growth within the blood-cells result in the destruction of the latter, a very serious thing to the patient if the organisms be at all numerous. If the spores be sucked up by an Anopheles, they undergo a complex change, and ultimately reproduce an incredible number of minute spores or ‘blasts,’ each capable of infecting man again if it can but win entrance into his body.

Under normal circumstances, for each Filaria larva which enters a mosquito, one Filaria issues forth, longer, it is true, and more highly developed, but not much changed. The malaria-parasite undergoes, in its passage through the body of the Anopheles, many and varied phases of its life-history. As the Frenchman said of the pork, which goes into one end of the machine in the Chicago meat factories as live pig, and comes out at the other in the form of sausages, ‘Il est diablement changé en route.’ The mosquito is as truly a host of the malarial parasite as man, and is as necessary for its full development as is man. Judging by the number and extent of the lesions in the insect’s body, it must suffer far more than man, and it is undoubtedly killed at times, and perhaps fairly frequently, by the parasite.

Whoever has watched under a lens the process of ‘biting’ as carried on by a mosquito, must have observed the fleshy proboscis (labium) terminating in a couple of lobes. The labium is grooved like a gutter, and in the groove lie five piercing stylets, and a second groove, or labrum. It is along this labrum that the blood is sucked. Between the paired lobes of the labium, and guided by them (as a billiard cue may be guided by two fingers), a bundle of five extremely fine stylets sinks slowly through the epidermis, cutting into the skin as easily as a paper-knife into a soft cheese. Four of these stylets are toothed, but the single median one is shaped like a two-edged sword. Along its centre, where it is thickest, runs an extremely minute groove, only visible under a high power of the microscope. Down this groove flows the saliva, charged with the spores or blasts of the malaria-causing parasite. Through this minute groove has flowed the fluid which, it is no exaggeration to say, has changed the face of continents, and profoundly affected the fate of nations.

It is an interesting fact that, amongst the Culicidæ, it is the female alone that bites. The mouth-parts of the male are weaker, and seem unable to pierce the skin. It has been suggested that a meal of blood is necessary for the development of the eggs; but the evidence for this is not conclusive. There must be millions and millions of mosquitoes in sparsely inhabited or uninhabited districts, in Africa, in Finland, in Northern Asia, and America, which never have a chance of sucking blood; and it is impossible to believe that these millions do not lay eggs.

The female is undoubtedly greedy. If undisturbed, she simply gorges herself until every joint of her chitinous armour is stretched to the cracking-point. At times even, like Baron Munchausen’s horse after his adventure with the portcullis, what she takes in at one end runs out at the other. But she never ceases sucking. The great majority of individuals, however, can never taste blood, and subsist mainly on vegetable juices. In captivity they cannot last longer than five days without food and drink; but they can be kept alive for weeks on a diet of bananas, pineapples, and other juicy fruits.

Anopheles is often conveyed great distances by the wind, or in railway trains or ships; but of itself it does not fly far; about five or six hundred yards—some authorities place it much lower—is its limit. Beyond this distance they do not voluntarily stray from their breeding-places. Both Anopheles and Culex lay their eggs, as is well known, in standing water, and here three out of the four stages in their life-history—the egg, the larva, and the pupa—are passed through. The larva and the pupa hang on to the surface-film of the water by means of certain suspensory hairs, and by their breathing apparatus. Anything which prevents the breathing tubes reaching the air ensures the death of the larva and pupa. Hence the use of paraffin on the pools or breeding-places. It, or any other oily fluid, spreads as a thin layer over the surface of the pools and puddles, and clogs the respiratory pores, and the larvæ or pupæ soon die of suffocation.

In Ismailia the disease has been reduced to an amazing extent, and quite recently remarkable results have followed the use of these preventive measures at Port Swettenham, in the Federated Malay States. Within two months of the opening of the port in 1902, 41 out of 49 of the Government quarters were infected, and 118 out of 196 Government servants were ill. Now, after filling up all pools and cleaning the jungle, no single officer has suffered from malaria since July, 1904, and the number of cases amongst the children fell from 34·8 to 0·77 per cent. The only melancholy feature about this wonderful alleviation of suffering due to the untiring efforts of the District Surgeon, Dr. Malcolm Watson, is that his fees for attending malarial cases have dropped to zero.

Thus a considerable degree of success has attended the efforts of the sanitary authorities, largely at the instigation of Major Ross, all over the world, to diminish the mosquito plague. It is, of course, equally important to try and destroy the parasite in man by means of quinine. This is, however, a matter of very great difficulty. In Africa and in the East nearly all native children are infected with malaria, though they suffer little, and gradually acquire a high degree of immunity. Still, they are always a source of infection; and Europeans living in malarious districts should always place their dwellings to the windward of the native settlements. Knowing the cause, we can now guard against malaria; mosquito-nets and wire windows and doors are a sufficient check on the access of Anopheles to man. If they could only be kept permanently apart, we might hope for the disappearance of the parasite from our fauna. In relieving man from the pest, all lovers of animals will rejoice that we are also relieving the probably far more acute sufferings of one of the most delicate and beautiful insects that we know.

Another elegant little gnat, Stegomyia fasciata, closely allied to Culex, with which, until recently, it was placed, is the cause of the spread of that most fatal of epidemic diseases, the yellow fever. Like the Culex, but unlike the Anopheles, Stegomyia has a humpbacked outline, and its larva has a long respiratory tube at an angle to its body, from which it hangs suspended from the surface-film of its watery home. It is a very widely distributed creature; it girdles the earth between the Tropics, and is said to live well on shipboard. It breeds in almost any standing fresh water, provided it be not brackish. The female is said to be most active during the warmer hours of the day, from noon till three or so, and in some of the West Indies it is known as the ‘day-mosquito.’

The organism which causes yellow fever has yet to be found. It seems that it is not a bacterium, and that it lives in the blood of man. It evidently passes through a definite series of changes in the mosquito, for freshly infected mosquitoes do not at once convey the disease. After biting an infected person, it takes twelve days for the unknown organism to develop in the Stegomyia before it is ready for a change of host. The mosquitoes are then capable of inoculating man with the disease for nearly two months. The period during which a man may infect the mosquito, should it bite him, is far shorter, and extends only over the first three days of the illness.

Very careful search has hitherto failed to reveal the presence of the parasite of yellow fever. By its works alone can it be judged. It seems that, like the germ of rinderpest and of foot-and-mouth disease, it is ultra-microscopic, and our highest lenses fail to resolve it. From the course of the disease and the nature of its host, it will probably prove to be something like the organism which causes malaria. The means of warring against Anopheles and Culex are equally applicable in the case of Stegomyia, but, since the last-named flies by day, they are more difficult to carry out, and more irksome to endure. By the intelligent application of these preventive measures the Americans have freed Havana for the first time from yellow fever, and have materially reduced the amount of malaria, and they have been equally successful at Panama.

King Solomon sent to Tarshish for gold and silver, ivory, and apes and peacocks, and at the present day people mostly go to Africa for gold, diamonds, ivory, and game. These are the baits that draw them in. Of the great obstacles, however, which have for generations succeeded in keeping that great continent, except at the fringes, comparatively free from immigrants, three—and these by no means the least important—are insignificant members of the order Diptera. We have considered the case of Culex and Anopheles; the third fly we have now to do with is the tsetse fly (Glossina), which communicates fatal diseases to man and to cattle and domesticated animals of all kinds.

There are at least seven species of the genus which received its name as long ago as 1830, when Wiedemann first described it. Perhaps the best known species is Glossina morsitans, which was named by Westwood.

The members of the genus Glossina are unattractive insects, a little larger than our common house-fly, with a sober brownish or brownish-grey coloration. When at rest the two wings are completely super-imposed, like the blades of a shut pair of scissors; and this feature readily serves to distinguish the genus from that of all other blood-sucking flies, and is of great use in discriminating between the tsetse and the somewhat nearly allied Stomoxys and Hæmatopota.

The tsetse flies rapidly and directly to the objects it seeks, and must have a keen sense of smell or sight, or both, making straight for its prey, and being most persistent in its attacks. The buzzing which it produces when flying is peculiar, and easily recognized again when once heard. After feeding, the fly emits a higher note, a fact recalling the observation of Dr. Nuttall and the present writer on the note of Anopheles, in which animal they observed that, ‘the larger the meal, the higher the note.’ The tsetse does not settle lightly and imperceptibly on the sufferer as the Culicidæ do, nor does it alight slowly and circumspectly after the manner of the house-flies, but it comes down with a bump, square on its legs. Like the mosquito, the tsetse is greedy, and sucks voraciously. The abdomen becomes almost spherical, and of a crimson red, and in the course of a few seconds the fly has exchanged the meagre proportions of a Don Quixote for the ampler circumference of a Sancho Panza. There is a good deal of discrepancy between the reports of the various sufferers as to the pain of the bite. No doubt different persons are very differently affected, and suffer to very varying degrees. Unlike so many of the blood-sucking Diptera, in which the habit is confined to the females, both sexes of Glossina attack warm-blooded creatures.

The fly always seems to choose a very inaccessible portion of the body to operate on—between the shoulders in man, or on the back and belly in cattle and horses; even inside the nostrils in the latter, or on the forehead in dogs. According to Lieutenant-Colonel D. Bruce, R.A.M.C., to whom we owe so much of our knowledge of this fly and its evil work, the female does not lay eggs, but is viviparous, and produces a large active yellow larva, which immediately crawls away to some secluded crevice, and straightway turns into a hard, black pupa, from which the imago emerges in some six weeks. Thus two stages, the egg and the larva, both peculiarly liable to destruction in the Culicidæ, are practically skipped in the tsetse—at any rate, in some species. On the other hand, this advantage is probably to a great extent counterbalanced by the smallness of the number of the larvæ produced, compared with the number of the eggs laid by the oviparous Diptera.

The genera of the Culicidæ which we have considered are found practically all over the world, but the genus Glossina, except that it just reaches Arabia, is fortunately confined to Africa. From the admirable map of the geographical distribution of the fly compiled by Mr. Austen we gather that its northern limit corresponds with a line drawn from the Gambia, through Lake Chad to Somaliland, somewhere about the 13th parallel of north latitude. Its southern limit is about on a level with the northern limit of Zululand. The tsetse, of course, is not found everywhere within this area, and, though it has probably escaped observation in many districts, it seems clear that it is very sporadically distributed. Mr. Austen further thinks that it may occur outside the boundary above laid down, and suggests that the great mortality amongst the horses in the Abyssinian campaign against King Theodore may have been caused by it.

Even where the tsetse is found it is not uniformly distributed, but occurs in certain localities only. These form the much dreaded ‘fly-belts.’ The normal prey of the fly is undoubtedly the big game of Africa, including crocodiles, but they are not the only factor in its distribution; the nature of the land also plays a part. There are the usual discrepancies in the accounts of travellers, especially of African travellers, as to the exact localities the Glossina affects; but most writers agree that the tsetse is not found in the open veld. It must have cover. Warm, moist, steamy hollows, containing water and clothed with forest growth, are the haunts chosen. Even within the fly-belt there are oases, due, perhaps, to an absence of shrubs or trees, where no flies are.

The tsetse fly belongs to the family Muscidæ, the true flies, a very large family, which also includes our house-fly, blue-bottle fly, etc. These flies, unlike Anopheles and Culex, are day-flies, and begin to disappear at or about sunset, a fact noted centuries ago by Dante:

The practical disappearance as the temperature drops has enabled the South African traveller to traverse the fly-belts with impunity during the cooler hours of the night. At nightfall the tsetse seems to retire to rest amongst the shrubs and undergrowth, but, if the weather be warm, it may sit up late; and some experienced travellers refrain from entering a fly-belt, especially on a summer’s night, until the temperature has considerably fallen.

The sickness and death of the cattle bitten by the tsetse were formerly attributed to some specific poison secreted by the fly, and injected during the process of biting. It is now, largely owing to the researches of Colonel Bruce, known to be due to the inoculation of the beasts with a minute parasitic organism conveyed from host to host by the fly. The disease is known as ‘nagana,’ and the organism that causes it is a species of Trypanosoma, a flagellate Protozoon or unicellular organism, which moves by means of the lashing of a minute, whip-like process. Since Bruce’s researches a number of Trypanosomas have been found causing diseases in various parts of the world. Thus T. evansii causes the ‘surra’ disease of cattle, horses, and camels in India. T. equinum produces the ‘mal de caderas’ of the horse-ranches of South America, and T. equiperdum is responsible for the North African disease called by the French the ‘dourine.’ T. theileri causes the gall-sickness, and there are others. These parasites were first seen by Gruby, who named them in 1843, in the blood of a frog; they live, not as does the malaria parasite, in the blood-cells, but in the fluid of the blood. The particular species of Trypanosoma which causes nagana is Trypanosoma brucei, and it does not attack man, and some goats and donkeys seem also immune; but, with these exceptions, all domesticated animals suffer, and in a great percentage of cases the disease terminates in death. Just as the native children in Africa form the source of the supply of the malarial parasite without appearing to suffer much, so the big game of the country abound in Trypanosoma without appearing to be any the worse. They are, in Lankester’s phrase, ‘tolerant’ of the parasite, and a harmony between them and the parasite has been established, so that both live together without hurting one another. Under a more natural condition of things than at present obtains in South Africa, the big game formed the natural prey of the tsetse; and, indeed, so dependent is the fly on the antelopes, etc., that, in places where the game has been exterminated, the fly has also disappeared. It is from the big game that the disease has spread. In their bodies the harmful effect of the parasite has through countless generations become attenuated, but it leaps into full activity again as soon as the Trypanosoma wins its way into the body of any introduced cattle, horse, or domesticated animal. Whether the Trypanosoma does any harm to the fly, or whether it passes through any stages of its life-history in the body of the fly, is still a debatable point. Possibly it does not, and the proboscis of the fly acts then simply as an inoculating needle.

The Report of Colonel Bruce, which was issued three years ago, shows that the sleeping-sickness which devastates Central Africa, from the West Coast to the East, is also conveyed by a species of tsetse fly. Writing over a hundred years ago of Sierra Leone, Winterbottom mentions the disease. ‘The Africans,’ he says, ‘are very subject to a species of lethargy which they are very much afraid of, as it proves fatal in every instance.’ Early last century it was recorded in Brazil and the West Indies; and in all probability the deaths which our slave-owning ancestors used to attribute to a severe form of home-sickness, or even to a broken heart, were in reality caused by sleeping-sickness. The severity of the disease, which always terminates fatally, is shown by the fact that in a single island—Buvuma—the population has recently been reduced by it from 22,000 to 8,000, whilst whole districts have been almost depopulated. In one year the deaths in the region of Busoga reached a total of 20,000; and it is calculated that although the disease was only noticed in Uganda for the first time in 1901, that by the middle of 1904 100,000 people have been killed by it. The disease is caused by the presence of a second species of Trypanosoma in the blood and in the cerebro-spinal fluid. The existence of this parasite has now been proved in all the cases recently investigated. Apparently the Trypanosoma can live in the blood without doing much harm, and only when it reaches the cerebro-spinal canal does it set up the sleeping-sickness. It is also found in great numbers in the lymphatic glands, especially those of the neck, which in patients infected by the parasite are usually swollen and tender. From the similarity of the parasite to that causing the cattle disease of South Africa, the idea at once arose that the Trypanosoma was conveyed from man to man by a biting insect. Along the lake shores a species of tsetse (Glossina palpalis) abounds; and it was noticed that if the fly, having fed off a sleeping-sickness patient, bit a monkey, the monkey became infected. Further, flies which were captured in a sleeping-sickness district were also capable of conveying the disease to healthy monkeys. The proof that sleeping-sickness is due to a Trypanosoma known as T. gambiense present in the cerebro-spinal fluid of the patient, and that it is conveyed from man to man by Glossina palpalis, seems now complete. Fortunately, like its congener, G. palpalis is confined to certain districts. The knowledge of these, and of the habits of this species of fly, will suggest preventive measures; and the brilliant research of Colonel Bruce and his colleagues, Captain Grieg and Dr. Nabarro, may yet save the much-tried African continent from the most fatal of recent diseases.

Finally, we come to a last class of disease which is of the utmost interest to the agriculturist and settler, and yet at present is but little understood. These diseases are caused by various species of a Protozoon named Piroplasma, and the diseases may collectively be spoken of as piroplasmosis. When they are present in cattle they are spoken of in various parts of the world as Texas fever, tick fever, blackwater, redwater, and many other French, German, Italian, and Spanish names. Heartwater in sheep is a form of piroplasmosis. Horses also suffer, and the malignant jaundice or bilious fever, which makes it impossible to keep dogs in certain parts of this country, is also caused by a Piroplasma. Finally, under the name of Rocky Mountain fever, spotted or tick fever, the disease attacks man throughout the west half of the United States.

The organisms which cause the disease live for the most part in the red blood-corpuscles, but they are sometimes to be found in the plasma or liquid of the blood. Unfortunately, we know but little about the life-history of the Piroplasma, or of the various stages it passes through, but we do know how it is transmitted from animal to animal and from man to man.

We have seen that the carrier or ‘go-between’ in the case of the malaria is the mosquito, and in the case of the sleeping-sickness is the tsetse fly. The Piroplasma, however, is not conveyed from host to host by any insect, but by mites or ticks, members of the large group of Acarines, which include beside the mites the spiders, scorpions, harvestmen, and many others.

The ticks differ from the insect bearers of disease inasmuch as the tick that attacks an ox or a dog does not itself convey the disease, but it lays eggs—for I regret to say here, as with the Anopheles, it is the female only that bites—and from these eggs arises the generation which is infective, and which is capable of spreading the disease. The tick which conveys the Piroplasma from dog to dog is called Hæmophysalis leachi. The brilliant researches of Mr. Lounsbury have shown that even the young are not immediately capable of giving rise to the disease. The female tick gorges herself with blood, drops to the ground, and begins laying eggs. From these eggs small six-legged larvæ emerge. These larvæ, if they get a chance, attach themselves to a dog, gorge themselves, and after a couple of days fall off. If their mother was infected they nevertheless do not convey the parasite. After lying for a time upon the ground the larval tick casts its skin and becomes a nymph, a stage roughly corresponding with the chrysalis of a butterfly. This nymph, if it has luck, again attaches itself to the dog and has a meal, but it also fails to infect the dog. After a varying time it also drops to the ground, undergoes a metamorphosis, and gives rise to the eight-legged adult tick. Here at last we reach the infective stage; the adult tick is alone capable of giving the disease to the animal upon which she feeds, and then only when she is descended from a tick which has bitten an infected host. Think what a life-history this parasite has! Living in the blood-corpuscles of a dog, sucked up by an adult tick, passed through her body until it reaches an egg, laid with that egg, being present while the egg segments and slowly develops into the larva, living quiescent during the larval stage and the nymph stage, surviving the metamorphosis, and only leaping into activity when the adult stage is reached. This most remarkable story probably indicates that the Piroplasma undergoes a series of changes comparable to those of the malaria organism when it is inside the mosquito; what these stages are we do not at present know, but Dr. Nuttall and Mr. Smedley at Cambridge, and many other observers elsewhere, are at work on the problem, and soon we shall have more light.

With regard to bovine piroplasmosis, Koch, and others have distinguished redwater fever, which is conveyed by Rhipicephalus annulatus, and in Europe probably by Ixodes reduvius from the Rhodesian fever, which is conveyed by Rhipicephalus appendiculatus, and I regret to say by a species dedicated to myself, Rhipicephalus shipleyi.

The heartwater disease of sheep and goats is similarly conveyed by Amblyomma hebræum, the Bont tick, and many farmers accuse Ixodes pilosus of causing the well-known paralysis from which sheep suffer in the early autumn; and there are many others, diseases such as the chicken disease of Brazil, which is so fatal to poultry yards, and which is conveyed by the Argas persicus.

I will not weary you with more diseases. I think I have said enough to show that within the last few years a flood of light has been thrown upon diseases not only of man and his domestic animals, but upon such insignificant creatures as the mosquito and the tick. I have tried to show how these diseases interact, and how both hosts are absolutely essential to the disease. We can now to a great extent control these troubles; the old idea that there is something unhealthy in the climate of the Tropics is giving way to the idea that the unhealthiness is due to definite organisms conveyed into man by definite biting insects. We have at last, I think, an explanation of why Beelzebub was called the Lord of Flies.

THE DANGER OF FLIES

It was Linnæus who first described this insect and named it Musca domestica, and de Geer who, in the middle of the eighteenth century, first described its transformation. In 1834 Bouché described the larva of the insect as living in the dung of horses and fowls. In 1873 the well-known American entomologist, A. S. Packard, reinvestigated the question, and L. O. Howard has recently written on the subject. In our own country C. Gordon Hewitt is publishing a monograph on the house-fly, which will, when completed, fill a long-felt want. Packard noted that in the August of 1873 the house-fly was particularly abundant, especially in the neighbourhood of stables. He was able to observe the insects laying their ova in clumps containing some 120 eggs in the crevices of stable manure, ‘working their way down mostly out of sight.’ The eggs hatched in about twenty-four hours, but he noticed that those hatched in confinement required from five to ten hours longer, and that these larvæ when hatched were smaller than those hatched out in the open. The eggs are oval and cylindrical, one twenty-fifth to one-twentieth of an inch long and about one-hundredth of an inch wide, and of a dull, chalky-white colour.

The little larva has not been seen emerging from the egg-case, but probably, as in the case of the meat- or blow-fly, Musca vomitoria, the eggshell splits longitudinally and the maggot pushes its way out. The length of the newly-hatched larva in its first stage (or instar) is seven-hundredths of an inch, and it remains in this stage about twenty-four hours, when it casts its skin and appears as a larger maggot three-twentieths of an inch long. In this condition it remains from twenty-four to thirty-six hours. After a second moult the maggot attains the length of one-quarter of an inch, and in this stage it remains five or six days. During its life the larva moves actively about amongst its surroundings, eating up the decaying matter, but avoiding bits of straw and hay. There is some evidence to believe that, if pressed for food, larvæ may devour one another. After living altogether some five to seven days, the larva somewhat suddenly turns into a dark brown pupa or chrysalis. The transition takes place very rapidly—in the course of a few minutes—and the pupa remains enclosed in the last larval skin. After another period of five to seven days in normal circumstances the insect hatches out, at first running around with soft and baggy wings, which, however, soon stretch out, harden, and dry. It is worthy of note that whereas Howard found the complete metamorphosis to take ten days, and Packard from ten to fourteen days, in the cooler climate of Manchester Hewitt finds it takes from twenty to thirty days. The last named gives some interesting particulars as to the effect of the weather upon the rate of development. It is believed that many flies pass the winter in the pupa state; the adult fly also survives the cold weather hidden away in cracks and crevices, from which it may from time to time emerge when the sun shines warmly.

When the larvæ are reared in too dry manure, they attain only one-half their usual size. Too direct warmth and the absence of moisture and available semi-liquid food also tend to dwarf them.

A word may be said about the distribution of the insect. It is practically cosmopolitan. As Mr. Austen records:

It is carried all over the world in ships and trains, and seems to be equally at home in the high latitudes of Finmark or in the humid heat of Equatorial Brazil.

The diseases which flies convey from man to man—which rendered them by no means the least formidable of the plagues of Egypt, and fully justified Beelzebub’s title of the ‘Lord of Flies’—are for the most part conveyed mechanically. The proboscis acts as an inoculatory needle. No part of the life-history of the disease-causing organism must necessarily be carried on in the body of the fly; it is conveyed mechanically and without change from an infected to a healthy subject. The mouth parts can pick up the anthrax bacillus, and if the fly then alight upon a wounded surface it will set up woolsorter’s disease. It, together with the flea, is accused of transmitting the plague bacillus, not only from man to man, but from rat to man. Flies are active agents in disseminating cholera; and anyone who has watched them clustering around the inflamed eyes of the children in Egypt, or in Florida, will not readily acquit them of being the active agents in the spread of inflammatory ophthalmia or of ‘sore eye.’

It is worthy of note that after exhaustive experiments on the tsetse fly (Glossina palpalis), which conveys that most fatal of diseases, sleeping-sickness, Professor Minchin and his colleagues, Mr. Gray and Mr. Tulloch, have come to the conclusion that the Protozoon (Trypanosoma gambiense) which causes the disease does not—as might be expected—pass through certain stages of its life-history in the fly, but is mechanically conveyed upon the biting mouth parts of the insect. The deadly parasite is, indeed, so easily cleaned off these appendages that a single bite is sufficient to wipe them off. A tsetse fly which has bitten an infected person will set up the disease in the next person (or monkey) it bites; but the insertion of the proboscis, quick and instantaneous as it is, serves to clean it—to wipe off adhering trypanosomes, and if it now bite a second person (or monkey), it fails to convey the disease. This is a most important discovery, and contrary to what we should have expected; but our knowledge of the history of the genus Trypanosoma is still too small to justify generalization, difficult as it is to avoid it. The diseases which in our country are disseminated by flies are all bacterial and all mechanically conveyed.

In passing, it is worth recording that, contrary to the usual statement that tsetse flies are confined to the continent of Africa, Captain R. M. Carter[8] has recently brought some back from the Tabau River and from other localities in South Arabia. Mr. Newstead has recognized the specimens as belonging to the species Glossina tachinoides. It evidently does not live on big game here, since, except the gazelle, game is absent. The Bedouins say that it bites donkeys, horses, dogs, and man, but not camels or sheep. It is at times so troublesome as to force the natives to shift their camps.

The common house-fly has been known for some time to be an active agent in the dissemination of bacterial diseases. In intestinal disorders—such as cholera and enteric fevers, which are caused by micro-organisms, the flies convey the bacteria from the dejecta of the sick to the food of the healthy. In the recent war in South Africa they are described in the standing camps as dividing their activities ‘between the latrines and the men’s mess-tins and jam rations.’[9] In the Spanish-American War in Cuba, and in the South African War, and in several recent outbreaks of enteric fever in the British army in India, flies have been proved to be the carriers of the Bacillus typhosus. Dr. Veeder[10] writes:

Similar records come from the Boer camp at Diyatalawa in Ceylon. The bacilli are conveyed direct, just as they might be by an inoculating needle. They do not pass into the body of the fly, neither do they undergo any part of their life-history in its tissue.

Dr. Sandilands[11] has recently investigated outbreaks of epidemic diarrhœa. He points out that the prevalence of diarrhœa follows the earth’s temperature, and does not follow the temperature of the atmosphere. It is a well-known fact that this illness is more prevalent in the houses of the poor than in the mansions of the rich. As Dr. Newsholme, late Medical Officer of Health for Brighton, said:

Compared with cow’s milk, which nourishes a very numerous progeny of bacteria, the bacterial content of Nestlé’s milk is very low, according to Dr. Sandilands. In certain seasons the cow’s milk is exposed to temperatures which favour an enormous multiplication of bacteria, and yet it is not then a frequent source of diarrhœa—in fact, mere numbers have little or no influence on the incidence of the illness. The greater number of cases are due to infection conveyed from some patient in the near neighbourhood and conveyed mechanically by flies.

The great attraction of the sweetened condensed milk for flies to some extent explains the greater prevalence of infantile diarrhœa among children fed on this preparation.

As was stated above, one of the most remarkable features in the prevalence of infantile diarrhœa is that it follows the rise and fall of the earth’s temperature, and not that of the air. In the same way the number of house-flies does not reach its maximum with the first burst of hot weather. The prevalence of these insects follows rather than coincides with periods of great heat. The flies, in fact, lag behind the air temperature and persist for a time after the hot weather has ceased. In other words, the meteorological conditions associated with an increase or a diminution of the prevalence of diarrhœa exercise a similar influence on the prevalence of flies.

The transference of the Filaria bancrofti, whose presence in the human body in the adult stage is associated with various diseases of the lymphatics, the most pronounced of which is the terrible elephantiasis, is due to more than one species of gnat or mosquito. It is true that no one has ever seen the actual transference of the Filaria from the biting organs of the Culex, Anopheles, Panoplites, or Stegomyia into the human body, but the circumstantial evidence is so strong that on it any jury would convict. Noè and Grassi have demonstrated a similar mode of infection for the Filaria immitis, which exists in the adult stage in such incredible numbers in the cavity of the right side of the heart of dogs, especially in tropical and in sub-tropical countries, that it is difficult to see how the circulation can be maintained at all. It is therefore interesting to note that the proboscis of our common house-fly frequently harbours a larval nematode which has been described by Carter[12] under the name of Habronema muscæ; and again (if it be the same species) by Generali[13] under the name Nematodum sp. (?), and again by Piana,[14] who is inclined to think it is the larval form of Dispharagus nasutus (Rud.). What the further history of this parasite is we do not conclusively know, but, judging by analogy—and in the case of the grosser parasites it is not always wise to do that—the nematode probably develops in some higher animal which eats the fly. Piana brings forward a good deal of evidence that this is the domestic fowl.

Another parasite which attacks flies is the fungus or mould Empusa muscæ, whose growth is fatal to the insect. The hyphæ penetrate into the body, and as they grow weaken the fly until it is unable to lift a leg, but remains glued by its viscid feet to the object upon which it rests. The fungus spreads and radiates out in all directions, covering the fly as with a velvety pile, and giving off countless minute spores, which are blown away, to alight, if they are lucky, on a further victim.

I think enough has been said to prove that flies are a very real danger to our community. I have refrained from giving the appalling statistics of our infant mortality, partly because of the difficulty of discriminating between the claims of the flies and those of other agencies which affect the lives of our babies—e.g., the insurance companies which do a large trade in insuring infants. Legislation has not attempted to control the latter. Sanitation might do much to destroy the former. In well-administered towns slaughterhouses no longer ‘fill our butchers’ shops with large blue flies’; they have been replaced by abattoirs, under proper inspection. Stables should also be segregated or controlled. The practice of backing the mansions of Berkeley Square by stable yards should either be given up, or the manure-heaps in which the flies breed should be under cover so close as to prevent the access of the fly. A layer of lime spread over the manure effectively prevents the fly laying. Creolin, in its cheap commercial form, is also recommended, sprayed over the manure-heaps every two or three days. It not only deters flies from ovipositing, but should they succeed in doing so it kills the resulting larvæ.[15]

Ross has shown us how to clear Ismailia of malaria; the Americans have rid Havana, for the first time in a century, of yellow fever; the same could be done with flies, if only the people liked to have it so. The motorcar, with all its destruction of nervous tissue, its prevention of sleep, its danger to life and to limb, has one great merit—it affords no nidus for flies.

CAMBRIDGE