Bacot and Martin have shown that while infection may take place as the result of the faeces being rubbed into the wound made at the time of feeding yet the ordinary way is probably by regurgitation from the oesophagus of the flea at the time of feeding.
Upon taking in plague septicaemia blood the bacilli multiply about the site of the proventriculus as well as distending the oesophagus. This makes an obstruction to the entrance of the stomach resulting in starvation of the flea. This naturally makes the flea more voracious and in the ineffectual muscular efforts to take in blood, regurgitation of the contents of the oesophagus occurs, thereby infecting the person upon whom the flea is trying to feed. This obstruction apparently may be overcome in some way as the plague-infected flea does not necessarily die. Still from lack of sufficient fluid such fleas are liable to be killed off if the relative humidity is low, as in dry weather. Further investigations have shown that the proventricular plug may be channeled, but in such case valve action is lost and the contents of the stomach are regurgitated, thus making such a flea more dangerous than one with an obstructed proventriculus. Such fleas may continue infectious for 67 days.
Very interesting in this connection is the fact that Heiser found plague-infected fleas in the desk of a European at Manila who died of plague. A mummified rat was found in one of the drawers of this desk, from which successful animal inoculations for plague were made. Heiser notes that data would indicate that these fleas probably remained alive 2 weeks after the death of the rat which brought about their infection. Another very striking finding during the same Manila plague outbreak (1912-1913) was that bedbugs found on the sleeping mat of a human victim of plague showed bipolarly staining bacilli.
Bacot has carried on experiments in which fleas infected two months previously and kept in a cool place could transmit plague. This would indicate the danger from plague-infected fleas which had been held in material packed away in boxes.
Method of Spread.—The spread of plague epizootics among rats seems to be rather by the fierce brown sewer rat, Mus norvegicus. The more delicate black house rat, Mus rattus, usually receives its infection from the sewer rat. When the rat dies the fleas leave the dead body and seek a new host, preferably one similar to the one just abandoned. The sewer rat reaching the basement of houses and dying of plague is deserted by his fleas. These will attach themselves to the house rats which go from basement to roof of the house and later these dying are abandoned by the fleas which, in the absence of a rodent host, will feed on man and infect him.
The house rat is rarely found in Europe while in many parts of the tropics it is common and in close association with man. The fact that the sewer rat avoids the upper portions of houses probably explains the greater infrequency of plague epidemics in Europe where this rodent is common. In former ages when the house rat was prevalent in Europe we had great epidemics there. Mus (Rattus) norvegicus is of stout build with a blunt nose and small opaque ears which barely reach the eyes when laid forward. The tail is shorter than the length of the head and body together (89% of such length). With Mus rattus we have a delicately built rat with a slender head and sharp nose. The ears are translucent and large and reach beyond the middle of the eye when extended. The rather delicate tail is about 25% longer than the length of the head and body taken together.
Recently it has been found that a guinea pig set free in a house suspected of having plague fleas becomes infected if such fleas are present. The fleas would probably prefer the guinea pig to man and such a measure would in some degree be protective to man. It is however for the detection of plague infection that the measure is employed and the guinea pig is termed the “Plague barometer.” In Madras there is practically an absence of Mus norvegicus although Mus rattus is present in numbers and the comparative freedom of the city from plague is striking.
The principal rat flea of the Orient is Xenopsylla cheopis. This flea is without combs like Pulex irritans, the human flea, but is of a lighter color and has an ocular bristle near the upper margin of eye and two bristles posterior to the antennae. In Europe and the United States Ceratophyllus fasciatus is the common rat flea. Many other species of fleas transmit plague and it is also possible that the bedbug may play a part in spreading infections from man to man. Vergbitski has transmitted plague from man to the rat by infected bedbugs. In Siberia, a marmot, the tarabagan, is supposed to play the part of the rat in plague transmission. In California, the ground squirrel, Citellus beecheyi, has become infected and may transmit the disease by its flea, Ceratophyllus acutus. In the Tropics plague tends to prevail only at times when the temperature is between 10° and 30°C. It is the effect on the flea of cold weather which causes the disappearance of bubonic plague at such times. The bacillus of plague can withstand freezing temperatures. Sunlight and drying are the especially inimical factors for B. pestis. Dry seasons are inimical to the spread of plague and it is especially in very rainy seasons that epidemics rage.
Chronic Plague in Rats.—The Indian investigators have called attention to the existence of a chronic plague in rats. In this we have chronic buboes, areas of necrosis in spleen and abscesses of the spleen. It is chiefly in the spleen that the lesions occur, thus differing from the acute plague in rats above described. Of 27,699 M. norvegicus, examined in Bombay, O.57% showed signs of chronic plague.
In the necrotic material plague bacilli can be found in approximately one-half of these rats although frequently the bacilli are nonvirulent. It is possible that this chronic plague in rats may serve as the reservoir of infection which keeps up plague epizootics from year to year. Plague in India, according to White is less virulent now than formerly and this is attributed to a greater immunity of the rats.