THE LIFE HISTORY and COMPARATIVE INFESTATIONS of POLYPLAX SPINULOSA (BURMEISTER) on NORMAL and RIBOFLAVIN-DEFICIENT RATS : Prese
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THE LIFE HISTORY AND COMPARATIVE INFESTATIONS OF POLYPLAX SPINULOSA (BURMEISTER) ON NORMAL AND RIBOFLAVIN-DEFICIENT RATS dissertation : Presented In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy In the Graduate School of The Ohio State University 3y DEFIELD TROLLINGER HOLMES,, B. Sc., M; Sc. The Ohio State University 1958 Approved by BriLd: Adviser Department of Zoology and Entomology ACKNOWLEDGMENTS X would like to make special acknowledgment to my adviser, Dr. C. E. Venard, Department of Zoology and Entomology, The Ohio State University, for his understanding encouragement and oontlnued assistance and stimulation throughout this research. Thanks are also due to Dr. D. M. DeLong and Dr. F. W. Fisk, also of this department, for the contribution of materials when needed; to Mr. Fhelton Simmons of the Columbus Health Department for kindly contributing rats from various areas In Columbus; to Mr. William W. Barnes and Mr. Roger Meola for technical assistance. Finally, I wish to express sincere appreciation to my wife, Ophelia, for her patience and enoouragement throughout this research and for her aid In the preparation and typing of this material. 11 TABLE OF CONTENTS Pag© Introduction .............................................. 1 Historical Review and Taxonomy .................. .... 6 Materials and Methods .....................................14 Life History ........................... .22 Incubation Period of the E g g ....................... 23 First Stage Nymph ....................................26 Second Stage Nymph ................... 2 8 Third Stage Nymph ................................... 30 Adult Female ........................................ 33 Adult Male ...................... 35 Moulting . ......................................... 37 Comparative Infestations on Normal and Riboflavin-deficient R a t s ................. .38 D i s c u s s i o n ........... 41 Summary . 46 Literature Cited .......................................... 48 ill List of Figures Figure Page 1 - Normal and Rlboflavln-deflolent R a t s ..........15 2 - Rlboflavln-deflclent Rat (Mild Symptoms) . 20 3 - Egg . .......... 25 4 - First Xnstar Nymph ............................. 27 5 - Seoond Instar Nymph ............................ 29 6 - Third Instar Nymph ............................. 32 7 - Adult Female .................................... 34 8 - Adult Male . .................................36 Text Illustrations Table Page 1 - Mode of Infestation .............. 18 2 - Time Required for Hatching of E g g B ............ 24 3 - Life Cycle of Lice on Normal and Deficient R a t s ............. 31 4 - Areas of Infestation on Ho s t s ............... .39 lv INTRODUCTION It has been noticed by various Investigators that laboratory rats varied widely In the number of rat lloe Polyplax snlnulosa (Burmelster) which they harbored. It was found that the more heavily infested rats had been maintained on vitamln-deflolent diets. Gyorgy (1938) noticed that heavy infestations of lloe became apparent In about 20 per oent of the animals that were kept for a sufficient length of time (more than eight to ten weeks) on rlboflavln-deflolent diet. Searls and Snyder (1939) found that rats most heavily Infested had been maintained on a vitamin A deficient diet. Kartman (1942) also noted that rats maintained on a vitamin A deficient diet showed heavy Infestations, but he stated that diets deficient In riboflavin also Increase the size of Infestations and that no particular vitamin plays the singular role, or even a limiting role, In determining the resistance of rats to their lice. The relation of host nutrition to other species of ectoparasites has also been studied, especially by De Melllon and co-workers (1946-47) In South Africa. These workers reported that egg laying of Clmex leotularlus was drastically reduced and that a large proportion of the eggs were sterile when the bedbugs were fed on thlamln-deflcient rats. When the tick Ornlthodorus moubata was fed on thiamin- deficient rata, there was a decrease In the rate of growth and size of nymphs, a prolongation of the interval between eaoh moult and an additional moult before maturity. When these two speoles of parasites were fed on rlboflavln- deflolent rats, normal growth and reproduction took place In all cases. An experiment with cattle lice was reported by Matthysse (1946) In which he suggested that vitamins A and D In the diet of the host apparently had no relation to the economy of these parasites. Barger and Card (1943) theorized that when fowls are either qualitatively or quantitatively undernourished, they become more susceptible to the attacks of lloe and mites than under conditions of optimal nutrition. Life histories of the lice of mammals have been determined for only a few species. Some aspects of the biology of Fedloulus humanus have been worked out by Nuttall (1917) and Buxton (1940). According to Weber (1939), our main knowledge of the biology of lice, sucking and biting, Is largely confined to ovlposltlon, hatching, and the fact that there are three nymphal stages which resemble the adults in habits and general appearance. It has been ascertained that the eggs are fastened to hair or feathers, and It has been observed that In most Instances the eggs will not hatch If kept at a temperature much below the body temperature of the host. 2 The body louse of man la an exoeptlon to the rule In that the eggs are usually attached to seams of the clothing worn next to the skin. The duration of the egg-stage and pre adult life is known only for a few species, and it varies with the temperature of the host as well as with the species of the louse. Hopkins (1950), In a comprehensive study on the host-assoclations of the lice of mammals, suggests a general life history of the lice, biting and sucking, as follows: egg stage, one to three weeks; the nymphal life, about a week to 14 days; and the pre-ovlpoBltlon period, from one to three days, thus giving the average period from egg to egg at about three to four weeks, and the number of generations a year about 12 to 14. Florence (1921) observed that the life history of the hog louse Haematoplnus suls was 29 to 33 days from egg to egg, which Is summarized as follows: Time from laying to hatching of egg..13 to 15 days First moult occurred after 5 to 6 days Second moult ocourred after ............... 4 days Third moult occurred a f t e r .......... 4 to 5 days Sexual maturity occurred after ............ 3 days There Is also little Information on the longevity of the adults, but for several species It is shown that this period Is not great. Unfortunately, much of this Information has been obtained by keeping the lice under unnatural con ditions and therefore Is of little value. Watts (1918) gives the duration of a generation of the hog louse Haematoplnus as 30 to 40 days, of which half represents the adult stage. 3 Orauford and Benson (1941) stated that for the cattle louse Haematoplnus eurvsternus the maximum longevity of the adults was 16 days for the female and ten days for the male. In Pedloulus humanus and Phthlrlua nubls the average life of the adult is about one month. According to Bacot (1917) and Buxton (1940), In Pedloulus the difference In length of life between tbe sexeB 1b small. Matthysse (1944) found the maximum life of the female oattle-bltlng louse Damallnla bovIs to be 42 days. The number of eggs laid has been determined for a few species and varies according to species. Lamson (1917) states that the cattle louse Haematoplnus eurvsternus lays from 35 to 50 eggs. Parthenogenesis has not been reported for most species, but Matthysse (1944) found parthenogenesis to be normal in Damallnla bovls. Hopkins (1950) states that in theory a single fertilized female could give rise to an enormous infestation within a very short period of time, but he suggests that this Is unlikely to occur. Elohler (1940) records the rise of an artificial Infestation of chewing lice from 200 to 14,000 In 80 days. However, he stated that because of biological checks such a great lnorease does not generally occur. Buxton (1940) has shown that a biological check in the case of Pedloulus humanus is the Injury to females which Is Inflicted by the males while making numerous attempts to mate. This occurs in populations where males exceed females. 4 No complete life history of the Incubation period of the egg and the longevity of the nytnpha.1 stages and the adult stage was found for Polyplax aplnulosa. Thus the purpose of this writer's Investigation was to work out the complete life history of the rat louse Polvplax aplnulosa and to oompare the degree of Infestation on rats kept on rlboflavln-deflolent diet and on normal diet. ft HISTORICAL REVIEW AND TAXONOMY The origin of the sucking lloe Is not very clear, but It Is generally accepted that lice are descended from Psocid-llke ancestors, and they have In the past been placed In the superorder Psoooldes, a position which appears to express their relationship with the other Insects better than any other which has been suggested. The relation of lice to the Psooldae was perhaps first suggested by Packard (1887)- Kellogg (1896 and 1902) came to the same conclusion. Lice belong to the primitive hemlmetabolous Insects, In which group the newly hatched Insects resemble the adult In most respects. They have formerly been divided into two orders: the Mallophaga, or ohewlng lice, and the Anoplura, or sucking lice. The ordinal name to be applied to the lice Is In dispute. Leach (1815) gave the name Anoplura to the whole group, and Nltzsch (1818) named the chewing lice Mallophaga, The latter, together with Ryncophthlrlna (Ferris, 1931) for the elephant-louse, has never been seriously disputed, but the discovery that the chewing and sucking lice are closely related and should be regarded as suborders has created ohaos as to the use of the name Anoplura. A few authors use this name for the whole group (applying the name Slphunculata to the sucking lice), while the great majority continue to use It for the sucking lice alone, as was 6 universal until a few years ago.