LACTA'I‘ION STUDIES IN MICE. ' TECHNIQUE THESIS m THE DEGREE OF M, 8... Kenneth B. DeOme . ‘ 3934‘ IN 111 WWW 129 301093 0174 ~ 'n. .m- I' .1 "v _ 1‘.- :“I’ t ”11".V‘v. _.;: 3' t" ' 1).“ ’7‘: 2‘. v ‘ ‘ .. . .' f‘td‘uh? . -. ‘1. ...- n. I'- v 34". ‘ . I.‘ ‘2 ‘ ,1“: 4 ~! 531:4: ». .1 1 .. ‘.' ‘. I \- I pr «0 ‘ .‘ , I‘ ' ‘ b. , 'L 1 v a» g: I ‘ ." ‘-“_(.g‘v.‘nlv J , ‘ .I;:.. ...". :o f.“"‘.““;‘ a ‘ o “l‘.'.. n,.$."7,__ ..y ’ ”‘1 "I “1., 9' A3” '0} 4 1," ‘11.: . a ‘ ”I": 5;" ”554.13; . ' I" ‘ I n5,» ‘1 '.r.~ 11' . - 17:. .1 o?" -...v -‘ s ‘, h, . I. we ’1'} Q“ d ' “’5 I. V ‘_‘.3| 9: :7 ‘L x’ ’ - hv-‘§(£p ’7‘_ g; . '3, at ’f mM' ' , w‘: 7? . _. 9. .v. . W . h l .x- . , . 'I. J. -: .2" ‘ . 'L’ «3 J 'W 'A‘M’- ~ r0 .0 I {TWA I -"§ ...11 u . - ., . ' 'e . ~" ‘ "‘ "s“ ‘ “‘1‘ y . ., :.. 1 7, .1 r. . I01 ‘0 ‘ . _ v. V . 2 - _ : '._.| ". f .1 -‘ .'_ __ g. . 1 ' ”a. :* .2; w ‘. :3."Ptfi‘ ’1‘, LL.“‘ . J: x ...-w- . -- - -' ~ ‘ ' '44“) UV" 9’s '3‘ ' 7* V‘" .. .. . w w‘m fr“ ‘- 4 .. ‘IL'IV#&."“.ZV ‘ "I‘I... 'm (€- '1“ F- ‘3 \fi‘ '- 1' '{nr‘w’fl‘ ‘4 QC"! 3 '1. ' - .... r " ‘. r ': “(.{Z‘E‘E-VJ‘aN‘JW- " “- .‘ v.4: vi." . 0 - k J-’~ " -. u,’ J. , . ‘. _ . ,c ‘ r n" 1‘ " 6 V ‘ . W":- W I a’~‘d._, ‘ . _ ' . ,_ ' -. ' . >1 .. - <‘ _ .' - V . ,' ”.1 '5 . f“ ' ' U. _‘,L .. . ., "-3 . v.‘ ..2‘ '.., .. . ._ _ . . ,p' ‘t u LACTATION STUDIES IN MICE I. TECHNIQUE THESIS FOR THE DEGREE OF M. S. _ r" A Kenneth B¥‘DeOme * 1954 THE-lid LACTATION STUDIES IN MICE I. TECHNIQUE Thesis Submitted to the Faculty of the Michigan State College in Partial Fulfillment of the Requirements for the Degree of Master of Science by. Kenneth B. DeOme July, 1954 ACKNOWLEDGMENTS The author wishes to express his appreciation for the assistance offered by his associates in the course of the experiments described in this paper. To Dr. H.R. Hunt, who suggested the problem and under whose thoughtful supervision it was carried out, special gratitude is due. Mr. Leo Klever has been especially helpful in preparing the apparatus necessary. Part II III VI VII VIII 1? 29 5? 4? 59 6. 7? 89 99 10. 11. 12. TABLE OF CONTENTS ........................... ......................... ................................ .......... .................... The Time Required for Suckling ........... The Number of‘Young pa. Litter ........... Method of Adding‘Ioung'Mice_... ........... The Initial Growth Period ................ F°°d9 Nesting “at9r1342.9?§.W9???- 9?9????? Isolation of Females ..................... Other Factors to be Condidered ........... Errors due to Urination and Defecation ... The ROUtine Of 8. Lactation 000000000009000 The General Consideration of Lactation_.......... A Test Of the StandardTechnique ooeeoeeooeeeooee The Animals U386. coco...oooooooooooooooeeoooooooo 1. ............... ............................... ------------- .......... The Choice of Comparable Samples .......... CDCDQCflU'IH 10 11 11 12 12 12 13 14 15 17 19 20 21 26 26 Part IX XI XII XIII XIV XVII XVIII TABLE OF CONTENTS CONTINUED 2. Comparison of Fourteen Day Samples Results Obtained from Mice #60-#37 ............. The Effect of Temperature on the Experiment .... The Dailwaain in Weight of the Young Mice as an Indication of the Rate of Milk Production . The Effect of Oestrus upon Lactation ........... ..................... ................................. CODOlHSiODS oooooooooococo-000000000000000000000 Bibliography cooooooooooooooooooooooosoooooooooo ................................... .......................... ......................................... Plates coocoocooooooooooooooooooooooeeoooooooooo 27 30 32 35 37 41 45 46 48 74 96 -1- Part I INTRODUCTION The genetic basis of lactation has been investigated extensively for many years. The material available for such studies is largely confined to records gathered by cattle improvement associations. The advanced registry system.main- tained by the various dairy breed associations, and the dairy breed improvement association records have offered a great mass of data for biometrical studies on the inheritance of milk production. Such competent investigators as John 'd Goren, M. H. Fohrmmn, W.E. Agar, R. R. Graves, H.D. Goodale, and many others have used the data from these records as a means of attacking the problem. It must be borne in mind, however, that such breeding records were not compiled from controlled breeding experiments, but were merely performance records of selected animals togeth- er with records of their ancestry. This fact offers two very serious objections to their use in genetic investigations- namely, the records represent only a selected portion of the whole cattle population (those whose records for breeding and production make them eligible), and the records are from Itté‘ ings made to produce a certain type of cattle rather then to discover genetic relationships. In view of the limitations mentioned above, those investigating the inheritance of milk and butterfat product- ion have been confined to the use of the correlation technique. -2- Much.suggestive material has been derived by this method, however, and no attempt to minimize it's value will be made here. For instance, such studies have pointed out that the process of milk production is at least partly hereditary, as would be expected. John W. Gowen, working at the Maine Agric- ultural Experiment Station, has published a number of analyses which show that daughters of high producing cows tend to be high producers; also, that the daughters of certain bulls make high records, while the daughters of other bulls are usually poor producers, and that there is a higher correlation between the production of full sisters than half sisters, etc. Studies on correlation between production records and body conformation have also been.made with the result that pro- duction "at the pail" plus a knowledge of ancestry is the only reliable method available at present for selecting high producing animals. R.R. Graves ('26) states, "The evidence seems to point to both parents contributing equally to the inheritance governing the milk and butterfat producing capacity of their daughter', and again that, ”when the records of a large number of daughters are compared with the records of their dams, there is a limited correlation, or a tendency for high record daughters to come from high record dams“. H.D. Goodale ('27) calculated a sire's breeding index from the observations of Cole, Castle, and Gowen on crosses of breeds differing markedly in production levels. He found that the offspring from such crosses tended to be intermediate between the two parental types, though not -3- exactly half way between the two.‘ I. E. Agar ('26) pointed out that the correlation coefficient between mother and daughter for red poll cattle of Australia was .295 1' .071 for the total milk production and .255 I .060 for total butterfat production. 1!. H. Fohrman ('26) from a study of the entry and re-entry records of the Guernsey, Jersey, and Ayshire cattle concluded that the production of a cow depends about 60% upon heredity and about 40% upon environment. Such data, while inclusive, show that heredity plays a large part in the production of milk and butterfat in cattle. They give no idea, however, of the number of factors involved, nor of the mode of inheritance. It is extremely improbable that such information will ever be extracted from the records of breed associations. It is also improbable that any organization will soon undertake the task of carry- ing on controlled experiments among cattle to determine the exact genetic structure influencing milk production. The excessive cost of maintaining such a project, the small number of offspring produced by a dam, the length of time required to carry out the experMent, and the difficulty of obtaining breeding stock representing the whole cattle population would make such a venture difficult if not impossible. A' search for a method of over-coming these difficulties provided the motive for this study. It has been realised by a number of people that small rapidly breeding laboratory animals might be used to help solve the problem if some adequate method of measuring the -4- quantity of milk given by them.coudd be developed. In hmman mothers, Holt & Rowland ('22) were able to estimate the amount of milk given by weighing the infant before and after nursing. Enxman ('33) adapted this method to mice, obtaining quantitat- ive measurements of milk production by taking the difference between the weight of a litter before and after nursing. By this technique he was able to construct milk curves for albino mice. With this method developed it seemed possible to use muse as a means of studying the genetics of lactation. It will be the purpose of this paper to describe the technique used in studying lactation in mice, to study the milk production of mice in successive lactations, and to discuss some of the factors influencing the experiment in question. ‘5- Part II THE STANDARD TE’CHN IQUE The technique used in.measuring the amount of milk given by an individual mouse is a modification of that suggest- ed by Ensman ('53). The difference between the weight of a litter before and after nursing is due to the milk obtained by suckling. Since the mother's milk is the only possible source of food until the eyes of the young mice are open, the above assumption must be correct. A fairly accurate quanti- tative measurement of the amount of milk given can be obtain- ed, then, if the number of young in the litter is kept large enough to take all of the mother's milk. The technique des- cribed below is built about this idea, and has gradually grown through.many trials and failures. It may be necessary to modify it still further as new relationships are revealed by future work. THE CAGE It is evident that the mother must be kept separated from her young except at such times as measurements are being made. During the early stages of the experiment the mother was left in a regular stock cage and the young were kept in a tin can partially filled with paper shavings. This plan worked quite well but was abandoned since it was thought that the situation would be more nearly normal if the young mice could be kept within sight and hearing of their mother. Another plan.was tried whereby the young mice were -6- left with their mother for 6 hours‘and then separated automatically from her for the next 6 hours before measure- ments were made. This was accomplished by placing the mother and her young in a small cage of i inch hardware cloth. This small cage was suspended in a glass battery Jar by means of a fine wire which was attached to the end of a lever. The lever was fastened by means of a swivel clamp to a heavy ring stand. A 500 e.c. flask was suspended from the end of the lever opposite the cage containing the mice, in such a position that it would nearly counterbalance the weight of the cage and mice. A rubber tube connected the flask on the lever with a flask full of water which.was placed about one foot above the lever. A stop-cock was placed in the rubber tube which was fastened to the alarm winding stem of an alarm clock. The alarm clock was set at 2:00 o'clock. When the alarm ”went off”, the winding stem reversed, turning the stopcock to an open position and permitting the water to flow from the upper flask through the stop-cook into the flask on the end of the lever. The added weight of the water in the flask cverbalanced the weight of the small wire cage and.mice, thus causing them.to be raised up a few inches from the bottom of the battery jar in which.the cage was suspended. The young mice would then fall through the t inch hardware cloth into the batter jar, but the mother mouse, due to her greater size, was unable to follow theme The young were unable to suckle then until 8:00 o'clock when measurements were made. This plan, though good in theory, was abandoned, due to the un- natural conditions involved. -7- Small cages made of galvanised iron and hardware cloth were constructed, removing both of the objections mentioned above. One of these cages is shown in Plate I. These cages measured 5% inches X 8 inches x 9 inches. They were separated into two compartments by a partition of fine hardware cloth. Both compartments were accessible from a door in the front of the cage. A small door, which could be opened and closed from outside the cage, furnished commun- ication between the two compartments. The back compartment was provided with the same paper shavings, food, and water as was used in our regular stock cages. The mother mouse was kept in the back compartment under normal conditions. The front compartment, which was smaller that the back, was provided with enough paper shavings to make a nettf‘fior the young mice. lo food or water was placed in this compartment. The small cage permitted the young mice to be kept with their mother, separated from her by a fine hardware cloth partition only. Her interest in the young was maintained since she was constantly aware of their presence. This cage provided a much more natural arrangement than any other device developed to date. In addition it was compact, well ventilated, and minimised the chance of error in handling the young mice. The adult female mouse need not be handled during the feeding period since she soon learns to enter the front compartment when it's entrance door is open. THE TEMPERATURE Early in the course of the experiment it was found that the young mice must be kept warm when they were separated -3- from their mother. They were normally warmed by the heat of their mother's body and the structure of the nest built by her. They became very inactive when cold and consequently did not suckle normally. To overcome this difficulty, an incubator was constructed in which the temperature was main- tained at 28 degrees to 30 degrees C. The whole cage was placed in the incubator so that the mother was subjected to the same temperature as the young. Under these conditions the young mice were normally active. The exact influence of temperature will be discussed more fully later. THE SUCKLING PERIODS Normally , young mice suckle very frequently. No method has been devised as yet to measure the amount of milk obtained by the young under these conditions. An arbitrary time was adopted for the suckling period, based largely on the convenience of the experimenter. It was decided to make measurements at twelve hour intervals. The suckling periods were arranged at 8:00 o'clock A. I. and 8:00 o'clock P. l. At these two periods the young were weighed, allowed to suckle , and weighed again. The date, time of day, time required for suckling, weight before suckling, weight after suckling, the difference between the two weights, and the hunter of young in the litter were recorded at each meas- urement. THE TIME REQUIRED FOR SUCKLIIIG The young mice were normally hungry enough to suckle immediately when given the opportunity. The older litters will finish suckling within an hour. The younger litters require more time.than'this. After a certain time the mother will -9- 1eave her young. This is taken to“ mean that the young mice have taken all of her milk. However, it is possible that the mother might leave her young for a variety of other reasons. Since the only way that the operator can tell when the young mice have taken all of their mother's milk is by observing the time at which suckling ceases: it was decided to subject this to test. Female #9 normally suckled her young for about 50 minutes and then left them. If the young had taken all of the milk that female #9 produced during this period, then there should not have been any further gain in weight even if they had appeared to suckle again within the next hour, since very little more milk should be available. For eleven successive measurements the young were weighed as soon as their mother left them and the amount of milk calculated on that basis. The young were then returned to their mother, who would immediately nestle over them with every appearance of suckling. The young were allowed to remain with her for the remainder of a two hour period, and then weighed. In ten of the eleven trials the young showed a loss in weight during the second 'suckling' period,whilein one trial they showed aslight gain. From these trials it seems that the mother leaves her young usually when she has no more milk for them. An arbitrary limit of two hours has been established at which time the young were removed and weighed, whether they appeared to be nursing or not. The time required for suckling depended upon the ind- ividual mother and was, roughly, inversely proportional to the age of the young mice. -10- THE NUIBER OF YOUNG PER LITTER The number of young per litter was not held constant during the experiment, for it should be large enough to take all of the mother's milk. There is no direct way of telling whens mouse's mammary glands are empty, as in cattle. Some idea can be gained, however, from the rate at which the young mice gain in weight. The normal average daily increase in weight for young mice during the first 14 days after birth, as indicated by Robertson's ('il) work on growth, is about .5 of a gram. This figure represents litters of various sises and is probably a fair average where the young have all the milk that they need. In other words, a gain of .5 of a gram.per day per mouse would indicate that the young had all the milk they could eat. For a lactating female which produced large quantities of milk a gain of .5 of a gram per day would only indicate that the young had more milk available than they could take. Since there is no way of determining the rate of milk production of a mouse other than by the young, it was decided to keep the daily gain in weight much.lower than .5 of a gram. This was done by in- creasing the size of the litter to such a point that the young would gain less than .5 of a gram per day per young mouse. In many cases it was necessary to decrease the number of young in the litter rather than increase it, since the young would lose instead of gain and would finally become emaciated and die. An average of the daily gains for the experiment proves to be about .2 of a gram.per mouse per day. This is much less than the average daily gain where -11- an abundance of milk is available. Such a small increment assured that the young were hungry enough to take all of the milk that their mother had and yet gain enough to keep them active. It was thought best to start each female with a large litter so that she would produce the maximun amount of milk as early as possible. Small litters were, therefore, built up to seven as soon as possible after birth. These litters were then increased or decreased as the individual case de- mmnded. METHOD OF ADDING YOUNG MICE It is not difficult to add young mice to the already existing litter if a few precautions are taken. It is advis- ible to add mice of about the same size as those in the litter. A little I'pine oil" rubbed on the hands while handling the young to be added will usually cover up the odor of their former nest. The young mice should be introduced after the suckling period so that they have an opportunity to acquire the odor of the nest before the adult female finds them. K THE INITIAL GROWTH PERIOD Observations of the feeding habits of very young mice shows that they suckle aLmost continually. It was found, if the young are removed from their mother at birth and re- turned only at twelve hour intervals for nourishment, that they die or become very emaciated before a week has passed. The second lactation of females #30-#37 failed for this reason as will be indicated later. To overcome this it was decided to allow the young to remain in the nest with their mother for the first four days. This allowed them to gain strength enough to withstand the twelve hour feeding schedule during the course of the experiment. This four day period will be refered to hereafter as'the initial growth period'. FOOD; NESTING MATERIAL, AND WATER "Fox chow',1nanufactured by the Purina Milling Company, was used to feed the motherathroughout the experiment. This is the regular stock ration in this rodent colony and has proven to be a very well balanced diet. It is made in small bricketts which could be placed in the back compartment of the experimental cages. Food was always available. Water was furnished from.gravity bottles which were thrust through the tops of the cages. wood shavings were used on the floor of the cagesas litter. Paper shavings were found to be the most satisfactory nesting material. ISOLATION OF THE FEMALES The females were isolated in the small experimental cages and placed in the incubator at the temperature mentioned above. The small door between the two compartments was left open so that the female mouse would have access to both come partments. She would usually build her nest in the front compartment. By the tune the litter was born she had ample time to become thoroughly acclimated to the new situation. OTHER FACTORS TO BE CONSIDERED There is no doubt that psychological factors enter into the experiment. rho natural temperament of mice, like people, varies. A part of the task of the operator, then, is to know the individual mice with which he is working and to handle them accordingly. Some mice are very tame and do -13- not object to any type of handling while others must be handled with the greatest of care. A number have been found which can not be used even with the most thoughtful attention. Unnecss- ary noise or quick movements about the cages should be avoided. All of this is a matter of experience and cannot be adequately described. The whole procedure requires endless patience. It was found that some mice are particularly sensitive to the odor from the hands of the operator on the young mice. This can be easily overcome_by rubbing a little pine oil on the hands be- fore attempttng measurements. ERRORS DUE TO URINATION AND DEFECATION An unavoidable error entered into the experiment due to the uniration and defecation of the young mice during the suckling period. The mother mouse licked the young thoroughly while nursing, which evidently stimulated urination and defecation. She appeared to eat these waste products in the process of licking the young so that no quantitative este mate of their weight could be obtained. It is evident that the difference between the weight of a litter before and after suckling would be less than the weight of the milk consumed, by a quantity equal to the weight or the waste products. This error would be quite constant, and always in one direction since it would never tendto increase the differ- ence in weights but rather to decrease it. Such an error would not effect the results since only the relative ability of mice to lactate was being studied. It was, therefore, disregarded. -14- THE ROUTINE OF'A LACTATION It seemaadvisable to describe in detail the method of carrying out the process outlined above. The female mice to be tested were bred and then isolated in the experimental cages. These cages were then placed in the incubator as described. The small door between the compartments was left open so that the adult mouse could build her nest in the front compartment. The cage was then inspected each day and the date of birth and initial number of young were recorded. The sise of the litter was raised to seven if smaller than that number. Twelve hours before the initial growth period was over, the small door between the two compartments was closed so that the mother could not suckle the young. On the fifth day the young were removed from.the cage and weighed at the regular time. The gate between the two compartments was opened and the mother would usually enter the front compartment without further persuasion. The gate was then locked shut and the young returned to the front compartment with their mother. As soon as suckling was completed, or at the end of two hours as the case may no, the gate was opened again and the mother driven into the back compartment. The gate was locked shut again to prevent her return. The young were then removed and weighed again. The records were written up at this time, as described above. Any adjustment in the size of the litter, revealed by a study of the gain in weight of the young mice. was mmde before returning the young. They were then returned to the front compartment until the next suckling period. This process is repeated for each.mouse every twelve hours for the duration of the test. The precautions mentioned above must be observed throughout the experiment. -15- Part'III THE GENERAL CONSIDERATION OF LACTATION Studies of the rate of milk production have been confined largely to cattle. The author has been unable to find any conclusive quantitative studies on milk production in any other animal, excepting the recent paper by Ensman('55) dealing with albino mice. A number of authors have tried to estimate the relative rate of milk secretion in such animals as rats, mice, and rabits. All of these studies were made in connection with studies of the growth of the young. Robertson ('74) in his growth studies assumed that the differing growth.rates of litters of mice of the same size must be due to differences in the amounts of milk given by their mothers. in cattle, Brody, Ragsdale, and Turner ('25) fouend that the curve of the production of milk rose grad- ually until the thirty-fifth day, onthe average, and then declined gradually until the end of the lactation period. They foudd a close correlation between the character of the rise of the curve and the curve of monomolecular reaction in chemistry. The period of decline in the curve they found to be similar to the curve expressing the formula of recurrent reactions. Weatherfordaflsed the method of Golgi, which correlates the cytological picture of the cell with the rate of secretion. He found that in mice the quantity of milk produced increased until the tenth day and then gradually decreased. His work does not indicate what the total length of the lactation period might be. McDowell, Gates, and ~16- McDowell ('29), and Parks ('26) and Enzman and Pincus ('32) have assumed that the rate of milk production in the small rodents could be calculated from.the rate of growth of the litters of young mice conlnming the milk. The validity of this assumption will be considered later. It will suffice to say here that such a method does not take into consideration the natural differences in the ability of various nursing animals to utilize food. Such differences would be expressed in the rate of growth of individuals on the same diet and under exactly the same conditions. Turner correlated the general morphology of the whole mammary gland with the rate of milk production. By making whole mounts of the mammary glands of lactating mice he was able to study the number of lobules present in the entire gland. He assumed that the number of lobules present was proportional to the rate of mdlk secretion. His results agree with those of weatherford. .17- Part IV A TEST OF THE STANDARD TECHNIQUE Enxman ('33) was able to obtain quantitative meas- urements of the mdlk of albino mice. His results were obtained by the use of a technique much.like that described above. His method differed from.mine in that he allowed the young mice to eat solid food as soon as they would. Thus, the young were weaned during the third week. His records were taken over a period of twentybone days. During the third week it was necessary to correct his gross measurements for the amount of solid matter eaten by the young mice. Such a procedure introduces unnecessary complications, which the author belives are overcome in the technique described above. The type of curve obtained by hnxman agrees quite well, for the 14 days, in which.mice receive only milk, with those to be presented in the following discussion. The problem.to be discussed in the remainder of this paper must be clear at the outset. It is the desire of the author to devise a technique that is reasonably accurate and then to test that technique on a group of mice. It's applica- tion over a number of successive bactation of the same mice should demonstrate the reliability of the method. Gowen has shown that the amount of milk produced by a cow in- creases until she is about eight years old, or the average. Shmilar variations would be expected in the successive lact- ations of mice, so that it would be unreasonable to expect a mouse to produce the same quantity of milk in her first, second, third, and fourth lactation. However, if a group of females are considered, the same order of yields should be -18- expected over successive lactations, providing that the method used regulates the environmental factors closely enough. But the conditions effecting the lactation of mice are not well known, so that we could not expect too much from any technique until all of the environmental factors can be controlled. . _ Therefore, if the conditions of the external en- vironment are carefully controlled, and the data collected by the use of the technique mentioned should place the mem- bers of a group of mice in the same relative order with reference to the amountof milk produced, then the technique used would be proven sufficiently accurate. Data will be presented in the following pages to show that our method meets these requirements. _. W No attempt was made in this experiment to meas- ure the maximum.milk production of mice. Our object was to determine production under the specific conditions of the investigation. The external environmental factors were kept as constant as possible throughout the trials. No breeding experiments ofjgenetic nature have been carried out to date. It is hoped that the invention of a.well proven technique will make it possible to discover the genetic mechanism of lactation in mice. -19- Part V THE ANIMALS USED The animals used in the course of the experiment are fully described in Table # I. The bulk of the data to be presented was obtained from the twenty animals listed. Some additional remarks will be mmde on the first lactations of other animals not indicated in the table.“ These particular mice were selected on thebasis of their_availability at the beginning of the experiment,with no peggeived idea of their preformance.‘_Each animal was ear marked so that it could always be identified. _ Females #1-#12 inclusive were treated as one group, females #30-#57 asa second group. Measurements were made on all of the members of a group at the same time, so that the conditions of the experiment would be as nearly alike as possible. Part VI A RESUME' OF THE EXPERIMENT The first, second, and fourth lactations of group #l§#12 were-carried out strictly according to the standard technique. During the third lactation of these females a temporary incubator was construcEd by enclosing a section of the shelves with a sheet of heavy canvas. Two large electric light blubs were placed in this enclosure to supply the nec- essary heat. While the third lactation was still in progress, the outside temperature became high enough to permit the open- ing of the doors and windows of the colony house during the day. No thermostatic control was provided for the temporary incubator and the temperature fluctuated within wide limits. This arrangement provided extra spacebut proved to be a poor substitute since it was subject to great variations in temp- erature. The results obtained during the third lactation will be considered later. The first and fourth.lactations of females #SO-#37 were carried out as required by the standard technique. During the second lactation an attempt was made to shorten the initial growth.period to two days, with the result that the young mice failed to suckle properly. The second lact- ation failed completely from this reason. The third lactation of group #39§#37 was carried out at room.temperature instead of 280-3000,, which proved to be too low and resulted in failure again. 7 . Since only the relative ability of mice to produce mdlk was being studied, no attempt was made to obtain maximun milk production. Special feeds were not used since the cond- itions were kept as nearly normal as possible. -21- Part VII LACTATION IN MICE Tables #2, #3, #4, and #10 represent the records of females #2-#12 inclusive for the first, second, third, and' fourth lactation: Each entry represents the sum of two meas- urements, one taken at eight o'clock in the morning and the other at eight o'clock in the evening. No corrections of any kind have been applied to the figures. In a few instances measurements were not obtained for reasons beyond the control of the Operator. Such.omissions in the records were estimated by taking the average of the four adjacent measurements from the record book. This is accepted as good practice in the keeping of dairy records. . _ . , _ The length of the individual records is quite variable. It was originally planned to continue the measurements over a period of only fourteen days. The reason for this decision will be considered later. In practice, however, the measure- ments were continued as long as the cages were not needed for the next series of mice. _The result of this practice was the accumulation of a number of complete lactation records. . no records were obtained for females #4, #5, #6, #7, #11, and #12 during the first lactation. Litters were_born to these femmles, but they did not suckle normally. No clue can.be gained frmm the records for these failures since the females in question did not lactate. .Females #4, #7, #11, and #12 were only three months old when introduced into the exp- eriment. Experience has shown that it is often impossible to use animals of this age because of their excitable nature. -22- Female #1 diedcduring the first lactation; Her records are not included. Female #4 died in the interval between the second and third lactations, therefore her re- cord is missing from the third and fourth lactations. Females #11 and #12 died during the interval between the third and fourth lactations and them records are not included in the fourth lactation. Figures #l-#lO represent graphically the production of tenfemales whose records are complete. _Digure #11 shows graphically the daily mean production for all. animals in each lactation. It must be remembered that measurements were not begun until the fifth day, so that’thevfirstday indicated on the graphs is the fifth day after partuition. A study of figure #11 will show that the average production starts on the fifth day with from one to two grams of milk. The rate of production rises rapidly until the max- imum is reached between the sixteenth. and nineteenth days. a maximum average daily production of 3.896 grams was reached on the sixteenth day of 3:119:13? lactation. From this peak production the curve slop? gradually downward to the endof lactation between the thirty-ninth and forth-fourth days. The production curves of individual mice show con- siderable variation from the mean. The variations in the production from'day to day are very striking. The general shapefiof‘the production curve, the time at which-the peak production is reached, the duration of lactation, and the total amount of milk produced, also show considerable variation. -23- lhile these daily variations are very striking, it is interesting to note that there are similarities between the successive records of certain mice. Complete lactation records are available for three females for two successive lactations. Figures #8 and #9 show the records ofnthe second and third lactations of female #11. The general shapes of the two curves are much alike. In the_second lactation the peak was reached on the twenty-first day, which was two. days later than the peak for the whole population in thesecond lactation. In the_third lactation the peak was, reached on the seventeenth day, which was_one day I later than the mean peak for the third lactation. Lactation ceased on the fortieth day for the sec- ondlactation and on the forty-third day‘for the third lactat- ion. The total amounts of milkwere“ 91.99 grams for the sec- ond lactation and 8...; grams for. th. third lactation. Theproduction‘cur'ves of female #8.er the second and third lactations, figures #5 and #6, showno similarity as to general shape and duration of lactation. The time at which the peak production was. reached was in‘each‘ case one day later than the mean of the group. The totalanounts .of milk produced were 45.59 grams for the second lactation and 45.26 grams for the third lactation. The records of the second and third lactations of female #5 do not show such similarities. \f .- f y , _ . . , Figure #2 showing the third lactation of female #6, and figure #10 showing the second lactation Of female #12 are of interest, since the general shape of the curves and the durations ofilactation are so very different. The curve for female #6 covers a period of 57 days but at no time exceeds the mean of series.#2-#l2 females in their third lactation by more than .54grams. The curve for female #12 is for only 24 days but exceeds the mean of females #2-#12 for their second lactation by 2.56 grams at'the highest point. .- The total amounts cf milk produceddby‘ the twofemales are very nearly equal, that of female #6 being 57.21 grams and that of female #12 57.51 grams. The production curve of female #10 for the second lactation, figure #7, is of'the same general shape as that ‘of the mean for that lactation. The complete production curves are of interest since they show the entire course of the process of lactation. Young mice used in the experiment were kept dependen'j‘ on their mother's milk for nourishment so that she would produce milk as long as she was able. The extreme variability of that part of the production curve after the twentieth day is to be ex- pected since neither natural nor artifical selection has acted upon it. The duration of lactation in mice would be of the same nature since selection has acted only’upon that part of the curve during which the young normallyflsuckle. A study of the complete production curves will show that there are as number of marked depressions in the curve of each animal. These points of depression tend to recur at rather regular intervals. The rhythmical character of the curve suggests that cyclic influences might effect the product-. ion of milk. The nature of such forces will be discussed later. Considering it' s size, the mouse shows a remarkable ability toproduce milk. Female #11 gave 6.56 grams of milk in one day. At the age of three months she weighed 51.61 grams e -25- a daily production of 6.56 grams represents 20.1% of her body weight. An inspection of the records will show that daily records of more than four grams are very frequently obtained. Enzman records a few daily records of more than 7.00 grams. This is not surprising when one considers that the weight of the suckling mice usually exceeds that of the mother before the nursing period is over. i The recordslpresented agree very well withlthose given by Enzman ('55). His records are, as a whole, slightly higher. He made measurements at sixhour intervals, which might account for the increase in production. ‘Observations on the suckling habits oflyoung mice. indicate that they nurse yeryfrequently. It is possible that “the tetallamount of milk produced by a mouse lactating under normal conditions would be muchhigher thanthat indicated by the records by Enzman and ourselves. This is known to be the case in cattle, and milkings are made more than twice a day when the highest possible amounts of milkuare desired. In this experiment it was not thought necessary to make measurementskmore often since onlyjth‘e relative ability of_m_ice to lactate__was_ to be 9124,1951? , . . , From the records shown, it seems that the amount of milk produced by mice is a measurable‘quantity. This being so, there is no reason why mice can not be used~ in a genetic study of lactation. The greatest difficulty lies in under- standing and controlling the external and internal environ- mental influences affecting the process of lactation. Part VIII A STUDY OF SUCCESSIVE LACTATIONS THE CHOICE OF COMPARABLE SAMPLES The length of time required to obtain complete lactation records for a large number of animals over a number of lactations, has made it advisable to consider the reliability of comparable lactation samples. The seven day yield, properly timed and supervised, has been ' used with much success in cattle. It seemed reasonable to adapt some similar scheme here.‘ _ g The first fourteen days of_measurement,_that is, from.the5th918£h.days after partuition were selected as a sample period for the following reasons; __ __ _ 1. During this period the young mice are normally dependent upon their mother's milk fornourishment. _This period would,then, be free from.any complications due to feeding on solid food whilestill nursing. ' __ _ _ " ' f 2. This sample covers_the time of rise in product- ion. The peak is reached between the sixteenth and nine- teenth days. 3. This period is a fair sample of the whole lac- tation. If the total milk production of the animals, whose complete lactation records are shown in Figure:#i to #10 are arranged in order of decreasing quantities of mdlk pro- duced, this order will correspond exactly to the order of their fourteenth day samples, with the exception of female #4. This is shown in_Table # XIV. -27- COMPARISON OF FOURTEEN DAY SAMPLES FOR SUCCESSIVE LACTATIONS A test of the adequacy of the standard technique (previously described) to measure genetic differences between animals, would be to determine whether under constant environ- mental conditions a group of females maintained their relative positions in successive lactations. Data will be presented to show that the method describe in the section on technique actually reveals these inherent differences. A number of the animals used failed during one or more of the lactations. As a result, the records of only five animals are complete for four successive lactations. The data for these five are shown in Table # VI, where they are arranged in the order of decreasing quantities of milk produced during each lactation. The order of the animals in Table #.VI is constant for the first, second, and fourth lactations with the exception of the position of females #3 and #10, which have changed . places from the first to second lactation. These three lactat- ions were carried out under the exact conditions of the stand- ard technique. The order in the third lactation is altered considerable. The possible reasons for this change of order will be discussed under the heading of "The Effect of Temp-. erature on the Experiment". It should be said here, however, that the third lactation was not carried out strictly under the conditions specified in the standard technique. _Even under the altered situation in the temporary incubator, the changed order of the five animals is explainable on the basis of temperature variations within the incubator. A study of the five females whose records are com- plete for four successive lactations shows that these mice produced their greatest quantities of milk during their second lactation. The table below shows the mean daily production of each of these five mice for four successive lactations.c The, data is taken from the fourteen day samples of their records. Lactat 16;; 9.42 A}: age 349 am First- 1.950 2.012 1.528 5.407 6.257 Second 2.102 3.038 1.848 5.664 2.706 Third 2.105 1.214 1.126 1.408 6.088 Fourth 1. 662 2.462 1. 406 2. 901 2.106 This table shows that mice #2, #3, #8, and #9 produc- ed their greatest quantities ofmilk during the second lactation. ‘0“39.#10 produced the greatest amount during the first lact- ation. . _ A , The rate of milk production varies in mice in success- ive lactations. While the number of animals considered is too small tobe_conclusive, it appears that the greatest quantity of milk is produced in the second lactation. Gowen has shown that cows produce their greatest quantity of milk at 8.25 years of age,on.the average. . Figures #12 -#21 show graphically the fourteen day samples of females #2-#12 for their first, second, third, and fourth lactations. Several of the records are missing for the reasons indicated in Table # I and in the text. The shapes of the curves produced on successive lactations by a mouse are quite differentwhen only the fourteen day samples are consider- ed. The daily variations are very great but no variations of a cyclical nature are observed.“ It is evident frol.these curves that the daily fluctuations are not due to pecularities -29.. in the individual mouse which occur in the same fashion in successive lactations. It is interesting to note, in spite of the extreme variations, that the total quantity of milk produced during the fourteen day sampling period places the mice in the same order over successive lactations. This is shown in Table #VI.. . _ . _ The females of the #3o.#37 series show similar results. The first lactation of these animals is shown in Table # XI. The second lactation failed due to the reduced initial growth.period. The third lactation failed because the temperature of the room.was too low. The fourth lacta- tion included only fsmales #50 and #52. The others having died or failedto breed. The 14 day samples of females #50_ and #52 are shown below for the first and fourth.lactations, which were carried out under comparable conditions. gemale Eirst Lactation .Fougtg Lactation . 193221. £2.92 2233.1. 3.4223 #60 27.82 1.987 (grams) 2.6.61 1 .896 #52 26.17 1.666 _ (grams) a . 24.56 1.754 . This table shows, even after the failure of the second and third lactations, that an.animal will preform in same manner in successive lactations if the conditions of the standard technique are followed. The agreement between the first and todrflzlactations is very close. _ . The data presented above, though.limited, shows that the technique described controlls the external environmental conditions well enough to permit constant preformance over successive lactations. -30- Part IX RESULTS OBTAINED FROM MICE #50-#57 __ _ . Table # Xi; shows the records of females #50-#57 in- clusive during their first lactation.c The history of these animals is shown in Table #I. They were subjected to the usual conditions of the experiment during the first lactation with.very satisfactory results. . The temperature was maintained at 28°-50° C. in the incubator and the initial growth period was maintained at four days. _ , During their second lactation the initial growth period was reduced to two days instead of the customary four day period. All other conditions were kept the same as in the first lact- ation. {As a result of the reduced initial growth period, the young mice became very emaciated and dxfnot suckle well. The results obtained were very unreliable and measurements were attempted for only 7 days. During the third lactation of females #50-#57 the conditions of the experiment were maintained as usual, ex- cept that the experimental cages were kept at room.temperat- ure. The temperature of the rodent colony is thermostatically maintained at about 74° 1?. (262° 0.). Some small variation may take place due to the opening of doors and windows. _As a result of the reduced temperature the young mice became emaciated and failed to suckle properly. .. . . _ A _ unfortunately, only the records of females #50 and #52 were available for the fourth lactation. The other members having died or failed to breed as indicated in Table # I. .31- The fourth lactation was carried ont under the standard cpnditions with.an initial growth period of 4 days and a temperature of 28°-50° C. The records of females #50 and #52 are given in Table # XII. They compare very favorable with performance in the first lactation as is hown in Table # VI. - A l H H a . The data obtained from.females #50-#57 are of special value since they show that the standard conditions described on page 5' must be closely adhered to, if reliable results are to be obtained. They prove that the technique is fundamentally sound and that a.mouse will preform in essentially the same way over successive lactations if the conditions of the experiment are kept uniform as described under ”Technique". Part I THE EFFECT OF TEMPERATURE OF THE EXPERIMENT The conditions of the standard technique, described in the section above, specify a temperature of 28°~50° C. for nursing mice. During the course of the work, deviations from this standard have taken place. The results obtained under these variations in temperature are very interesting since they point out the necessity of maintaining the conditions as specified. .. _ _ _ .. . . .The third lactation of females #50-#57, inclusive, was carried out at room temperature which.is maintained automat- 16a11y. at about 74° 1?. (262° 0.). some slight variation in the room temperature doubtless took place. It was pointed out in the preceding section that the third lactation of females #50- #57 failed. Since mice in the regular stock cages at room temperature lactate sufficiently to raise normal litters, it could hardly be assumed that the reduction in temperature effected the lactating females. The young mice, however, were kept away from their mothers for eleven hours at a time so that they receive no heat from her body. They felt cold when handled and their movements were very sluggish, so that no doubt this sluggishness preventethheir nursing properly when the mother was introduced. The young are not protected much.by hair until about the tenth day. The third lactation of females #2-#12 was carried out in an improvised incubator which provided more adequate space. The necessity of accurate control of temperature was not appreciated fully until this lactation was finished. -33- This lactation was begun about April tenth and continued until lay twenty-fifth. During this time the weather became warm enough so that the steam could be turned off and the windows left cpen part of the time. The temperature of the room was ~then affected by the outside temperature. The incubator was heated by a large electric light bulb which.worked very well when the room temperature was constant, but which.offered no means of compensating for variations in the room.temperature. The exact range of temperature variation in the improvised incubator is not known since the lowest temperatures came during the night. Observation during the day showed variations from 26° 0. to 62° 0. Table # IV and # V show the results obtained from the third lactation of females #2-#12. Table #‘VI shows that females #2 and #10 maintained a production similar to the first and second laction, while_females #9, #3' and #8 showed marked decreases in production. The cages of females #10 and #2 were very close to the electric bulb which.was used to heat the temporary incubator. This doubtless explains their failure to show decreases in production. Females #9, #5, and #8, whose gages were some distance from the source of heat, show decreases. It is interesting to notice, how— ever, that the females showing decreases in production, fall into the same order in Table #‘VI as in the first and second lactations, but are lower in the series. Female #10 pro- duced more than female #2 in the first and second lactations. This is also true of the third lactation, although their. positions with reference to the other animals is changed. -54- It may be concluded that temperature is an important factor in the technique described. Deviations from the standard temperature of 28°-50° c. havetbeen shown to produce fluctations in the amount of milk obtained. It is probable that the reduced temperature effects the suckling mice rather than the lactating females by chilling them.until they do not nurse properly. -35- Part XI THE DAILY GAIN IN 'EIGHT OF THE YOUNG MICE AS AN INDICATION OF THE RATE OF MILK PRODUCTION A number of investigators, MacDowell, Gates and MacDowell ('29), and Entman anvaincus ('52) have assumed + that the daily gain in weight of the young mice might be taken as an indication of the amount of milk produced by their mother. The studies refered to hbove were not made directly upon the lactation of mice. -_ _ If such a relationship existed, there should be a constant relationship between the total gain in weight of theyoung mice and the total amount of milk consumed. This would be true only if the total number of days con- sideredeere the same in each case and_if the number of young in the litters considered were constant. Or expressed mathematically-- The total gain in weight of the young mice _ The total amount of milk consumed_ would equal a constant (k) if the assumptions were correct. The available records are tabulated in Table # XIII. Only litters in which seven young were carried through 14 con- secutive measurements are included. The weight of the young at the first weighing varied but little. The value (k) varies within wide limits, showing that no constant relationship exists between the gain in weight of the young and the amount of milk consumed. This variation is suggestive of a difference in the ability of young mice to turn food consumed into tissue. The records of females_#l2 and #10‘ show this clearly. Female #12 produced 57.21 grams of milk in 14 days. Her‘7 young gained 15.01 grams. Female #10 produced 57.91 grams of milk in 14 days and her young gained only 9.77 grams. -37- Part XII THE EFFECT OF OESTRUS UPON LACTATION Figures #l-#10 represent the record of one complete lactation for each of ten females. These curves show depress- ions which tended to recur at more orlless regular intervals after the peak production was reached. Figure #4 illustrates this point particularly well. The peak was reached on the sixteenth day, which was followed by a gradual decline until the nineteenth day. A sharp drop in productionntook place on the twentieth and twenty-first days, followed by a sharp rise. Another depression appeared on thecthirtieth and thirty-first days, and on thetthirty-ninth days. “Lactation stopped on the forty-second day. .The interval between these recurring periods Ofdepression was about ten days, which suggests that a cyclic influence may be the cause. a Figure #9Vshows similar recurring depressions on the t‘Irenty-second, thirty-third, and fourtieth days. The regular 10w points are not so clearly shown in the remaining figures though similar tendencies can be made. out in most cases. In eache of theseten figures a low point is reached at some time between the twentieth and thirtieth days, which 18 followed and frequently preceded by other similar depressions. _ . Except in the case of Figure #2, the highest point 111 aeurve is followed by a sharp decline within a very few dtays. 7 . The cyclic nature of these depressions suggests that they might be correlated with the oestrous cycle. Allen (‘22) -33- studied the oestrous cycle of normal mice and found that the stages of the cycle could be detected by daily microscopical examinations of the vaginal contents. Parks ('26) applied Allenfis method to lactating mice. He found, when the young were weaned normally, that oestrus recurred "at about the third week", but if lactation was continued by giving the female a second litter, oestrus reappeared from the twenty- fifth to the fortieth days. loClandish ('26) showed that the quantity of milk produced by cattle decreases during the period of heat. Allen made daily vaginal smears which he stained with eosin and hemotoxylin. Briefly his observations revealed the following periods in oestrus. . A Di-oestrust-Epithelium.(chiefly nucleated) and leu- ooytes are present during this stage. The vaginal smear tends to be stringy. . . Pro-oestruss-Lightly staining nucleated epithelial cells predominate the picture. During this stage the vaginal contents are serous in nature. . Oestrust-Non-nucleated, eosinophilous, cornified cells constitute the smears during this stage. As this period begins, the vaginal epithelium may be_dry; subsequently the contents of the vagina become granular. l Meta-oestrusz- The oestrus period is followed by a large infiltration of leucocytes which gradually removes the cornified elements. During this stage the vaginal content becomes first pasty and the milky. -59- During the fourth lactation of females #2-#12 daily vaginal smears were made and stained with eosin and hematoxylin. Plate # II shows the typical difoestrous smear with its scatter- ed polymorphonuclear leucocytes, moderate quantities of mucus, and epithelial cells in various stages of degeneration. A pro-oestrous smear is presented in Plate # III which shows the absence of polymorphonuclear leucocytes, and numerous lightly staining epithelial cells.‘ The typical oestrous smear is shown in Plate # IV containing nothing but cornified, heavily eosin0phil,non-nucleated cells. Plate #’V is firfimra\meta- oestrous smear. In this case polymorphonuclear leucocytes are very numerous in the later stagesbut scarce immediately following oestrus, while the epithelial cellsduring the early stages are almost entirely cornified, non-nucleated, and eosinophil. _ . . . . Plates #IV , #VII, #VIII, #VI, #IX, and #X show the first oestrus smears of females #8, #5, #5, #2, #9, and #10 repectively. Figure #21 shows the milk production curves and Table #X the records of these mice during their fourth lactation. In Figure #21 the days of oestrus are marked by small circles. Oestrus occured in these animals from the twenty-fourth to the thirty-first days. In each case the oestrus followed shortly after a high point in the curve, and occured at a low point. These figures agree very well with those given by Parkes, and according to him, oestrus should recurr again within seven days, on the average. The evidence presented here suggests that after the twenty-fourth days, the recurring oestrus is correlated with the periodic depressions in the milk curve. This may explain -40.- the cyclic variations seen in Figures #I-#X. Many other daily variations are present which cannot be explained now, unless they are normal for mice. . The figures given above were obtained from a study of 250 stained vaginal smears, from.females #2, #5, #5, #8, #9, and#10 and from a number of unmated females. _ It is interesting to note in the cases of females #9, #5,and #5, that the decline in the quantity of milk produced preceded the oestrous period. The forces which cause decreased milk secretion, then, must be separate from.those causing oestrus, for_otherwise the two events would always occur si- multaneously. It is probable that both oestrus and the decreased mammary activity are the results of a series of events in- volving the anterior pituitary gland and the ovary. -41- Part XIII DISCUSSION The purpose of this work was to develop a technique for measuring the milk production of mice and then to apply that technique to a number of animals over successive lac- tations. The procedure described in the section on "Technique" outlines the method used. H- The application of the standard technique has pro- duced the results discussed above. .It is regretable that the number of animals used was not larger but the time and equip- ment necessary to carry out the experiment limits the number of animals that it is possible to handle. .The method des- cribedmust be followed closely. Two measurements must be made 212;: day at the prescribed hours if success is to be attained. _ . .1 . . The results of_Enzman's_$f55) experiments are comparable to those presented above,only in,a limited way. He was studying the amount of milk giyen by a mouse during a normal lactation when the young were permitted to eat solid foodas soon as possible._ Hedid'not attempt to study the complete nor the successive lactations of mice.. The curves presented by him.are, when smoothed, of the first order. The production of milk rises gradually to the tenth day on the average and then declines to zero at about the twentieth day. . V The results of the technique described above show a somewhat different type of curve, since the whole lactation is considered. The production of milk rises rather sharply -42- to the l6th or the 19th day and then declines slowly to zero at some point between the twenty-seventh and fortieth days, depending upon the persistency of lactation. The peak pro- duction is reached somewhat later in this experiment than in Enzman's work. The reason for this difference is not evident. The young mice would need the greatest amount of milk at about the 15th to the 20th.days when their eyes are opening and be- fore they take enough solid food to satisfy their needs. The results obtained from these experiments show that the peak production is reached_during that period. _ _. The daily variations in milk production are very great in the results described above and by Enzman. Other than oestrus, no reason can be given for these variations at present. It may be that such variations are normal events in the lactation of mice, or they may be due to some environ- mental factors which are not properly controlled by the standard technique. Enzman presents several very smooth curves for females suckling only four mice. It is possible, however, that these curves represent the maximum.rate at which.the young mice con- sume milk rather than the rate at which the female could pro- duce it. Since the average size of litter is between 7 and 8 young mice, it is possible that the females producing these 9') smooth curves were capable of secreting more milk than four young mice could consume. The curves in this case would re- present the rate of milk consumption of four young when an abundance of milk was available. This consideration emphasizes the necessity of regulating the size of litter as is described 3% inqgection on "Technique". -45- The use of 14 day samples for comparing the product- ion of mice is equally applicable to Enzman's data, since his curves are of the first order. Such.a.method of sampling makes it unnecessary to carry animals through the complete cycle of lactation to determine their relative worth. The method of sampling is surprisingly accurate and very con- venient. _ ._ - It may be desirable at some future date to study the persistency of lactation in mice. _The technique des- cribed will make such a study possible._ Interesting variations in persistency will be noticed in the complete production records presented in Figures #I-#X. .Among these ten animals the persistency of_lactationvaries from 27 to 45 days, with as average of 56.9_days for the group. _ The mean daily production amongthese mice, as estimated from.the 14 day test, ranged from .665 grams . (female #50, first lactation) to 5.664 grams (female #9, second lactation). Similar variations can be seen in the duration of lactation, the time at which peak production is reached, and the general shape of the curve produced. If the technique is sufficiently accurate these variations may be looked upon as individual differences. .Consideration of the Figures #I'. #1 would tend to substantiate this view, as mentioned before. The available data shows that the greatest quantity of milk isproduced during the second lactation._ _ The recurrence of oestrus after pdrtuition has been shown to exert a cyclic influence upon the quantity of milk produced. It is doubtful whether’oestrus itself effects the mammary gland in such a way as to cause a decrease in milk secretion, but rather both events are the result of a series of influences involvingthe anterior pituitary gland, and the ovary.” This view is substantiated by the records of females #9, #5, and #5 as shown in Figure #XXI. In these cases the decline in milk production preceded the actual occurance of oestrus by several days showing that the force which.oaused the decrease in milk preceded that which caused oestrus. _ A The technique described and tested, should be sufficiently reliable to permit the use of mice in future studies on lactation. The application of this technique may make it possible to establish a situation among mice similar in every desirable feature to that in cattle. The rapidity with which.mice reproduce, the large_size_of litters, the ease with which they can be raised and studied, should make such a type of experiment exceedingly helpful in studying the complicated phenomena of lactation. Finally, the author hopes to be able , by the use of the information presented, to work out the genetics of lactation. Much.information concerning the physiology of lactation and reproduction will have to be gathered before this study can be completed. The only aim.of this paper was to develop and study a means by which this future work may be accomplished. -45- Part XIV CONCLUSIONS 1. A technique has been presented by means of which the entire milk production of mice can.be determined. 2. By means of this technique several mice have been carried through four complete lactations. ‘ 5. It has been shown that a group of mice will pro- duce milk in the same relative quantities in successive lac- tations if the conditions of the external environment are controlled according to the standard technique. 4. Ten complete lactation curves are presented which show that lactation may be continued as long as 45 days. The persistency of lactation varies from.27 to 45 days with an average of 56.9 days for the group of animals. 5. The fourteen day sample has been developed and proved to be a fair sample of the entire lactation. 6. lice produce the greatest quantity during the second lactations.. ._ 1.- __ _. _ 7. The recurrence of oestrus after partuition has been shown to correlate with rhythmical depressions in the lactation curve. a _ a A 8. Oestrus appeared between the twenty-fourth and the thirty-first days. . , _ 9. In three cases the decline in milk secretion preceded oestrus by several days so that the influence, causing the depression in the curve must be separate from that causing oestrus. -46.. BIBLIOGRAPHY Agar. W.E., 1926 Heredity and milk production. An analysis of the inheritance of milk and butterfat product-I ion in red poll cattle. Victoria (Australia) Dept. of IAgric. Jour. 24, pages 1-9. . . Allen, Edgar, 1922 The oestrous cycle in the mouse. Am. Jour. . . of gnat. 50:297- 521. _ _ . .. Brody, 3., Ragsdale, A.C., and Turner, C.W., l924 The relation between the initial rise and the subsequent decline of milk secretion following partuition. Jour. of. _ .. Gen. Phys. 53541-545. -u.. . ‘ Enzman, E.V ., 1955 The milk production curves of albino mice. . _ i Anat. Record 65:645-668. W Enzman, E.V. and Pincus, G., 1952 The effect on lactating mice of injecting an extract of the urine of .1 pregnancy. IAm. Jour. of Phys. 105350. . Fohrman, 11.11., 1926 Official records as material forstudying inheritance of milk and butterfat production. Jour. . of Dairy Science 93286-292. _ Gates, I.H., 1925 Litter size, birth weight, and early growth . rate of mice (Mus musculus).Anat. Record 29:185- 195. Goodale, H.D., 1927 A sires breed index with.apecial reference .. . . to milk production. Am. Naturalist, 61.559-544. Gowen, John W., 1924 Milk secretion. The Williams and Wilkins . company, Baltimore. . Gowen, John W., 1926 Studied in milk secretion XVIII. Trans- mitting qualities of guernsey sires for milk yield} butterfat percentage, and butterfat. Maine Agric. Exp. Station Bulletin # 529, page 1-48. Graves, R.R., 1926 Transmitting ability of 25 Holstein- . Friesian sires.U.S. Dept. of Agric. Bulletin 1672. Holt and Rowland, 1922 Diseases of Infancy and Childbirth. I I(quoted from.Enzman) I MacDowell, E.C.,Gates,'I.H.,Iand MacDowell, 0.0., 1929 The influences of thquuantity of nutrition upon the growth of the suckling mouse. Jour. of Gen. Phys. l5:529-545.I McClandish, Andrew 0., 1926 The influences of the period of heat on milk production. Jour. of Dairy Sci. 9(1);65-67. Parke, A.S., 1926 Observations on the oestrous cycle of the albino mouse. Proceeding of the Royal Society of I ILondon, series B, 100: 151-199. IIII _ Robertson, T. B., l9l6 Growth.studies on mice. Jour. of Bio. . I __ IChem. I24: 547-585. . I I . I . Robertson, T. B., 1917 Growth.studies on.mice. Jour. of Bio. I Chem. 51:567-514. . . Turner, C.W., 1952 The mammary glands. page 544. Sex and internal secretion. Iilliams and Wilkins Co. weatherford, H.L., 1929 A cytological study of the mammary gland: Golgi apparatus, trophospongium and other cytoplasmic canaliculi, mitochondra. Am. Jour. of Anat. 44:199-282. Part XVI Table # I Animals used in the experiment Female Strain Age at Wt. in Remarks begin. of grams ' exp. #1 F1 from P1 5 mo. 29.12 died during first ‘ 'cross of ' ‘ ‘lactation #2 dark eyed 5 mo. 28.78 black ' ' #5 agonti 9 5 m0. 28013 X ‘ ‘ ‘ #4 chinchilla d! 5 mo. 27.14 died between second ' ' ‘and third lactation #5 " 5 mo. 150.58 #7 " 6 mo. 62.02 #8 " 6 mo. 60.40 #9 fl 5 mOc 29e15 #10 " 6 mo. 29.91 #11 " 5 mo. 51.61 Idied between second ' " and third lactation #12 " 5 mo. 50.25 Developed tumor. killed between second and third lactation #50 dark eyed 4 mo.' 54.10 white #51 " 4 mo. 28.20 failed to breed for ' ‘ fourth lactation #52 " 4 mo. 28.45 #55 dark eyed 5 mo. 25.57 failed to breed for black ' ' fourth lactation. #54 dark eyed 5 mo. 28.92 failed to breed for ' black agouti ' ' 2 nd& 53d lactation. #55 pink eyed 5 mo. 29.17 developed infection dilute brown in ear. Killed during fourth lactation. #56 dark eyed 4 mo. 27.18 died during third white ' lactation. #57 ” 4 mo. 29.16 died during third lactation. -49- Table # II Daily production records of females # 2-#12 for the first lactation. Daze 32 9395 91514 915 9446 m e 300 Table# I 5 .46 .76 F.* F.* 6.5 6.. 6 .66 .29 7 .76 .29 8 1.07 1.15 9 1.79 1.16 10 1.24 1.50 11 1.00 1.62 12 1.52 2.10 15 1.05 1.29 14 2.62 5.90 15 5.46 5.10 16 6.76 5.64 17 5.65 5.25 16 5.75 5.51 19 2.96 2.92 20 5.27 5.25 21 22 25 24 25' 26561 55.26 54.26 Moan 2.06 2.14 -50- Table #II Continued Daily production records of females #2-#12 for the first lactation. 4 566 Tab1o#1 M6 ‘ 2449 £10 g£11 ' 94512 E. can 5 .59 1.51 2.26 F.* 6.4 1.064 6 .06 1.55 2.50 .956 7 .46 2.04 2.66 1.250 6 .54 2.26 2.75 1.550 9 .60 4.46 5.06 2.556 10 1.15 2.76 5.64 2.076 11 1.55 5.04 5.66 2.264 12 1.65 4.65 2.99 2.576 15 2.16 4.69 5.65 2.606 14 2.54 4.57 5.06 5.542 15 2.46 5.95 5.45 5.292 16 2.62 4.64 4.62 5.696 17 2.41 4.26 5.90 5.494 16 2.56 4.56 4.14 5.660 19 5.25 4.55 5.05 5.552 20 5.44 5.41 1.62 5.454 21 5.16 2.64 .57 2.125 22 1.65 1.66 1.655 25 2.22 5.66 2.950 24 2&,. 26541 55.50 66.62 52.06 lean 1.85 5.51 5.06 second lactation. Rsxs___2££_» ?#9 955:1, 2#5 Qié 2i!... 5 1.64 1.46 2.59 1.60 2.06 1.42 6 .46 .99 1.17 .99 1.42 1.25 7 .61 1.45 2.17 .97 .72_ 1.55 6 1.66 2.51 2.14 1.01 2.52 1.04 9 .57 2.65 2.57 .66 2.62 .61 10 1.25 5.97 4.41 .56 2.59 1.55 11 2.76 4.59 4.07 .26 2.92 .81 12 2.90 2.95 2.65 .14 2.66 4.27 15 2.72 5.46 2.66 .40 2.15 5.71 14 2.64 5.61 .96 .62 2.20 5.94 15 5.14 4.22 2.51 1.66 5.14 5.19 16 5.56 4.29 2.66 1.96 5.56 2.26 17 2.90 5.96 2.65 .67 2.26 .46 16 2.60 2.26 2.14 1.55 2.72 .00 19 2.22 1.65 2.52 2.74 5.00 20 .75 2.50 5.15 5.95 21 5.10 1.96 2.66 1.77 22 2.16 1.75 .51 25 5.07 .56 1.66 24 1.66 1.59 1.62 -51- Table.#III Daily production records of females #2-#12 for the Table #III Continued second lactation. Daily production records of females #2-#12 for the Da 6 2 _gi5 9#4 Agfs 9f6 9g:__ 25 2.60 1.47 2.49 26 1.11 1.86 2.40 27 1.29 2.51 1.45 26 1.77 .46 .90 29 2.15 .15 .79 50 1.45 .51 .42 51 2.52 .41 .75 52 .57 .52 .98 55 .01 .41 .66 54 .00 .00 .54 55 .00 .00 56 57 56 59 40 T6561 56.12 44.56 55.77 57.25 44.50 26.26 lean 1.670 2.956 1.754 1.266 2.617 1.675 -53.. Table #111 Continued Daily production records of females #2-#12 for the second lactation. 0416 2£§ «igfs 9£10 9£11 9£1g £63; 5 2.59 2.52 2.25 1.77 1.61 1.955 6 .66 1.56 1.19 1.90 1.62 1.219 7 1.45 2.05 1.69 2.66 .47 1.466 6 2.65 2.55 1.45 2.59 1.40 1.919 9 2.74 2.52 2.02 5.46 1.04 1.696 10 2.55 2.71 2.56 4.05 2.61 2.526 11 1.91 4.65 5.64 5.56 2.67 2.916 12 1.21 5.56 5.75 4.46 5.05 5.069 15 1.45 4.92 2.56 5.41 2.90 2.774 14 .54 4.51 2.57 4.72 5.47 2.727 15 1.97 4.15 2.56 5.46 2.96 5.000 16 1.55 5.66 2.16 5.91 5.96 5.015 17 1.65 5.56 4.57 5.61 5.09 5.056 16 5.02 4.45 5.54 2.24 4.14 2.765 19 5.17 5.40 5.95 4.55 5.66 5.266 20 4.11 1.17 4.41 4.75 5.21 5.105 21 5.51 4.15 5.95 .62 2.965 22 5.65 4.07 5.60 2.51 2.607 25 2.56 2.95 2.02 2.65 2.256 24 1.56 .96 5.52 5.56 2.150 -54- Tablo #III Continued Daily production records of females #2-#12 for the second lactation. gays £1648 9&9 ?#10 2#11 9fl2 ' Mean 25 .00 2.57 2.75 1.56 1.920 26 .55 2.69 5.16 .55 1.702 27 .00 2.99 2.90 .00 1.562 26 .00 5.55 5.55 .00 1.454 29 .00 2.60 2.05 1.265 50 2.56 2.26 1.546 51 1.14 1.50 1.160 52 1.10 .74 .762 55 .55 .67 .465 54 .45 1.26 .416 55 .17 .76 .257 56 .00 .50 .15 57 .05 .05 33 .05 .05 59 .06 .06 4o .00 .00 Total 45.59 55.66 76.55 91.99 57.51 lean 1.625 5.492 2.454 2.555 2.596 Table # IV . 0611y production records of females #2-#12 for the third lactation. 93;. 9#2 955 9g; Qfs 2#6 5 1.41 .22 5.4 ‘ .70 .06 2.51 6 1.47 .70 .26 .69 2.64 7 1.52 1.05 1.04 1.25 1.95 6 1.06 1.64 .20 1.16 2.05 9 1.65 1.07 .67 1.15 2.62 10 2.72 1.64 2.15 1.52 1.80 11 2.94 1.56 2.55 1.65 1.69 12 2.16 1.27 1.92 2.74 1.97 15 1.56 .62 2.71 5.64 2.59 14 1.71 1.41 2.74 5.45 4.56 ' 15 2.72 1.69 1.99 2.95 .40 16 1.64 1.55 5.99 5.56 5.00 17 4.15 1.49 5.42 2.26 2.96 18 1.66 1.27 5.54 2.21 2.97 19 1.65 .60 5.19 2.66 2.62 20 2.24 1.99 .76 2.51 2.96 21 1.56 1.99 .19 2.65 2.06 22 2.95 2.44 .61 1.60 1.96 25 .66 2.27 1.55 2.00 1.14 24 1.75 2.56 1.67 1.74 1.64 25 1.50 2.26 1.86 1.99 5.54 4800 Table # I ‘52.. -55- Table # IV Continued Dily production records of females #2-#12 for the third lactation. 56;: fifz 255 9f4 9#5 9#6 'g£z__ 26 1.44 1.59 9.4 1.66 .41 5.47 27 2.06 1.85 2.22 1.55 2.90 26 2.65 .65 1.66 1.55 5.10 29 4.05 1.07 1.50 1.75 2.56 50 5.59 1.27 .56 2.16 2.77 51 5.75 1.77 .27 .06 52 4.27 2.59 .60 .45 55 4.15 5.01 .62 .76 54 2.76 .92 .45 55 4.20 1.50 1.55 56 2.55 1.56 1.65 57 1.65 1.75 .64 56 2.71 1.41 .67 59 .60 .16 4o .26 .00 41 .09 42 .00 Total 67.40 56.57 ‘ 54.69 57.21. 64.20 lean 2.555 1.725 1.597 1.654 2.469 4 See Table # I Table # IV Continued Daily production records of females #2-#12 for the third lactation. Dazs Qfa 9#9 g#10 9#11 9#12 Mean 5 .79' 2.66 .26 .50 .67 .940 6 .00 1.15 1.19 1.55 .57 .996 7 .54 .75 2.57 2.41 .69 1.551 6 .24 1.51 2.99 1.11 .94 1.291 9 .67 .51 5.22 2.52 .66 1.512 10 .44 ;1.15 4.25 2.50 2.12 2.027 11 .76 .61 4.26 2.79 2.45 2.126 12 .51 2.55 5.50 5.46 .91 2.061 15 .65 1.05 2.99 4.95 2.22 2.296 14 1.10 .79 5.75 4.27 2.55 2.591 15 1.65 1.65 5.42 5.90 .54 2.091 16 1.96 2.25 5.74 5.41 1.64 2.656 17 5.07 2.11 5.61 5.27 2.55 5.106 16 5.52 1.05 5.50 6.56 2.67 2.695 19 1.62 1.62 2.69 5.25 5.56 2.666 20 2.70 .10 1.97 5.46 4.15 2.464 21 1.51 2.51 1.56 2.92 2.47 1.942 22 1.55 5.11 .26 1.05 4.45 1.974 25 1.42 2.67 .05 5.05 1.77 1.656 24 2.40 5.75 .44 2.64 2.65 2.162 25 2.55 2.40 .66 2.51 2.66 2.175 -53- Table # IV Continued Daily production records of females #2-#12 for the third lactation. 2319 9#6 9#9 9510 _gf11 9512 Mean 26 1.54 1.67 1.01 2.52 5.41 1.652 27 1.52 5.72 1.57 2.67 5.66 2.576 26 1.17 1.62 1.97 2.69 4.20 2.162 29 1.22 1.25 1.75 2.55 4.52 2.176 50 .91 .65 1.47 2.26 2.99 1.665 51 1.61 1.12 1.09 1.41 2.55 1.565 52 1.22 .96 1.00 1.50 1.97 1.456 55 1.16 1.27 1.14 .00 5.54 1.565 54 .66 1.70 1.05 1.00 .674 55 .94 2.12 1.45 1.46 1.502 56 .95 2.51 1.57 1.05 1.120 57 .75 5.96 1.24 1.09 1.144 56 .56 1.62 1.54 .95 .926 59 .26 2.07 .27 .59 .599 40 .00 2.47 .56 .17 .550 41 .00 .94 .16 .121 42 .14 .21 .055 46 .00 .00 .000 Total 45.26 65.56. 70.26 69.56 69.91 lean 1.156 1.621 1.898 2.550 1.654 Table # V -59- Table showing the lean Daily Production of All Mice for the First, Second, Third, and Fourth Lactation. ggzg_ lst.Lact. 2nd.Lact. 5rd.Laot 455.5665. 5 1.064 1.955 .940 1.562 6 .956 1.219 .996 1.675 7 1.250 1.466 1.551 1.610 6 1.550 1.919 1.292 1.966 9 2.556 1.696 1.512 2.065 10 2.076 2.526 2.027 2.061 11 2.264 2.916 2.126 2.055 12 2.576 5.069 2.061 2.151 15 2.606 2.774 2.296 2.460 14 5.542 2.727 2.591 2.065 15 5.292 5.000 2.091 2.545 16 5.696 5.015 2.656 2.916 17 5.494 5.056 5.106 2.555 16 5.660 2.765 2.695 5.066 19 5.552 5.266 2.666 20 5.454 5.105 2.469 21 2.125 2.965 1.942 22 1.655 2.607 1.974 25 2.256 1.656 2.150 2.162 -50- Table # V.Continued Table showing the Mean Daily Production of All Mice for the First, Second, Third, and Pourth.Lactations. Days lst.Lact. 2nd.Laot. 5rd.Lact. 4th.Lact. 25 1.920 2.175 26 1.702 1.852 27 1.562 2.576 28 1.454 2.182 29 1.285 2.178 50 1.548 1.885 51 1.180 1.565 52 .782 1.458 55 .465 1.565 54 .416 .874 55 .257 1.502 56 .150 1.120 57 .050 1.144 58 .050 .928 59 .080 .599 40 .000 .550 41 .121 42 .055 43 .000 44 -51- Table.# VI Mean Daily production of those females of the #2-#l2 series those records are complete for four successive lactations, arranged in order of decreasing milk production. Fourteen day samples are considered in each lactation. Dirst Lact. 8econd Lact. Third Lact. Fourth Lact. .gf’ hean ?# hean ?# Dean _2£__ Mean 9 5.407 9 5.664 10 5.088 9 2.901 10 5.257 5 5.058 2 2.105 5 2.452 5 2.012 10 2.706 9 1.408 10 2.105 2 1.950 2 2.102 5 1.214 2 1.662 8 1.528 8 1.848 8 1.126 8 1.406 Daily mean production of females #50 and #52 for the first and fourth lactations. First Lact. Second Lact. .Third Lact. Fourth Lact. 2£_f Mean 9# Mean 50 1.987 F.* F.* 50 1.895 52 1.655 52 1.754 a See Text-page 30 -62- Table # VII Daily production records of females #2-#12 for the first lactation, 5-18 days inclusive. ‘ 2g 931:2 9,95 254 9%!5 91946 95917 5 .46 .78 p.* 9.9 r.* 9.9 6 .68 .29 7 .78 .29 8 1.07 1.15 9 1.79 1.18 10 1.24 1.58 11 1.00 1.82 12 1.52 2.10 15 1.05 1.29 14 2.82 5.90 .15 5.48 5.10 16 5.76 5.84 17 5.65 5.25 18 5.75 5.51 Total 27.05 28.18 '0“ 1.950 20012 ,5 See Table # I lactation, 5-18 days inclusive. -55- Table # VII Continued ~ Daily production records of females #2-#12 for the first 251; ?#8 9#9 9#10 '9#11 9#12 ‘Mean 5 .59 1.51 2.28 F.* F.* 1.084 6 .08 1.55 2.50 .956 7 .46 2.04 2.68 1.250 8 .54 2.26 2.75 1.550 9 .80 4.48 5.08 2.558 10 1.15 2.78 5.64 2.078 11 1.55 5.04 5.86 2.264 12 1.65 4.65 2.99 2.578 15 2.16 4.69 5.85 2.608 14 2.54 4.57 5.08 5.542 15 2.48 5.95 5.45 5.292 16 2.62 4.64 4.62 5.896 17 2.41 4.26 5.90 5.494 18 2.56 4.56 4.14 5.660 Total 21.59 48.70 46.60 lean 1.528 5.407 5.257 4 See Table # I lactation, 5-18 days inclusive. -64- Table # VIII Daily production records of females #2-#12 for the second Days 912 9f5 9&4 9i: 91546 3L 5 1.64 1.46 2.39 1.80 2.08 1.42 6 .48 .99 1.17 .99 1.42 1.23 7 .81 1.43 2.17 .97 .72 1.55 8 1.68 2.31 2.14 1.01 2.52 1.04 9 .37 2.83 2.57 .66 2.82 .81 10 1.23 3.97 3.41 .56 2.59 1.55 11 2.78 4.59 4.07 .26 2.92 .81 12 2.90 2.95 2.83 .14 2.86 4.27 13 2.72 3.46 2.86 .40 2.15 3.71 14 2.84 C3381 .98 .62 2.20 3.94 15 3.14 €4222 2.51 1.86 3.14 3.19 16 3.36 4.29 2.66 1.96 3.38 2.26 17 2.94 3.98 2.83 .67 3.28 .48 18 2.80 2.26 2.14 1.53 2.72 .00 20281 29.68 42.55 34.77 13.43 34.80 26.25 lean 2.120 3.038 2.482 .958 2.284 1.875 -64- Table # VIII Daily production records of females #2-#12 for the second lactation, 5-18 days inclusive. M ‘ 913 254 3,45 9,5 _g£7__ 5 1.64 1.46 2.59 1.80 2.08 1.42 6 .48 .99 1.17 .99 1.42 1.25 7 .81 1.45 2.17 .97 .72 1.55 8 1.68 2.51 2.14 1.01 2.52. 1.04 9 .57 2.85 2.57 .66 2.82 .81 10 1.25 5.97 5.41 .56 2.59 1.55 11 2.78 4.59 4.07 .26 2.92 .81 12 2.90 2.95 2.85 .14 2.86 4.27 15 2.72 5.46 2.86 .40 7 2.15 5.71 14 2.84 $5381 .98 .62 2.20 5.94 15 5.14 44922 2.51 1.86 5.14 5.19 16 5.56 4.29 2.66 1.96 5.58 2.26 17 2.94 5.98 2.85 .67 5.28 .48 18 2.80 2.26 2.14 1.55 2.72 .00 Total 29.68 42.55 54.77 15.45 54.80 26.25 lean 2.120 5.058 2.482 .958 2.284 1.875 -55- 7651. # 7111 Continued Daily production records of females #2-#12 for the second lactation, 5-18 days inclusive. Days 9#8 9£g .Qflo 9f}; 2£12 nean 5 2.59 2.52 2.25 1.77 1.81 1.955 6 .86 1.56 1.19 1.90 1.62 1.219 7 1.45 2.05 1.69 2.88 .47 1.468 8 2.84 2.55 1.45 2.59 1.40 1.919 9 2.74 2.52 2.02 5.48 1.04 1.896 10 2.55 2.71 2.58 4.05 2.61 2.526 11 1.91 4.85 5.64 5.58 2.67 2.916 12 1.21 5.56 5.75 4.48 5.05 5.089 15 1.45 4.92 2.56 5.41 2.90 2.774 14 .54 4.51 2.57 4.72 5.47 2.727 15 1.97 4.15 2.58 5.48 2.98 5.000 16 1.55 5.86 2.18 5.91 5.96 5.015 17 1.65 5.56 4.57 5.61 5.09 5.152 18 5.02 4.45 5.54 2.24 4.14 2.785 265.1 25.88 51.51 57.91 45.90 57.21 lean 1.848 5.664 2.706 5.278 2.800 Table # IX Daily production records of females #2-#l2 for the third lactation, 5-18 days inclusive. 64m 942 246 294 9.95 956 gfl_ 5 1.41 .22 F.* .70 .08 2.51 6 1.47 .70 .26 .69 2.84 7 1.52 1.05 1.04 1.25 1.95 8 1.08 1.64 .20 1.18 2.05 9 1.85 1.07 .87 1.15 2.62 10 2.72 1.64 2.15 1.52 1.80 11 2.94 1.58 2.55 1.85 1.69 12 2.16 1.27 1.92 2.74 1.97 15 1.58 .82 2.71 5.64 2.59 14 1.71 1.41 2.74 5.46 4.56 15 2.72 1.69 1.99 2.95 .40 16 1.64 1.55 5.99 5.58 5.00 17 4.15 1.49 5.42 2.28 2.95 18 1.86 1.27 5.54 2.21 2.97 Total 29.47 17.00 27.66 28.55 52.24 lean. 2.105 1.214 1.974 2.022 2.502 4 See Table # I lactation, 5-18 days inclusive. Table # Ix- Continued Daily production records of females #2-#12 for the third Dale 9f8 2§9 9&0 9f}; 91212 lean 5 .79 2.66 .26 .30 .67 .940 6 .00 1.13 1.19 1.33 .37 -.998 7 .34 .73 2.57 2.41 .69 1.331 8 .24 1.51 2.99 1.11 .94 1.291 9 .67 .51 3.22 2.52 .68' 1.512 10 .44 1.13 4.25 2.50 2.12 2.027 11 .76 .81 4.28 2.79 2.43 2.126 12 .51 2.35 3.30 3.48 .91 2.061 13 .63 1.05 2.99 4.95 2.22 2.298 14 1.10 .79 3.73 4.27 2.35 2.591 15 1.65 1.65 3.42 3.90 .54 2.091 16 1.96 2.25 3.74 3.41 1.84 2.656 17 3.07 2.11 3.81 5.27 2.55 3.106 18 3.52 1.03 3.50 6.36 2.87 2.893 T0581 15.78 19.71 43.25 43.60 21.38 lean 1.126 1.408 3.088 3.082 1.522 lactation, 5-18 days inclusive. -68- Table.# 1 Daily production records of females #2-#12 for the fourth 23;. 5’42 9545 53:3 955 956 921.. 5 1.28 1.25 F.& 5.04 1.4 3.. 6 .77 1.97 1.62 7 1.04 2.06 2.06 8 1.54 2.18 1.95 9 1.55 2.19 2.22 10 2.08 1.80 2.57 11 1.82 2.48 1.86 12 2.26 2.04 5.11 15 1.82 2.40 5.82 14 2.08 2.55 2.02 15 1.58 2.51 5.79 16 2.55 5.45 4.52 17 2.50 5.10 1.16 18 5.25 4.54 1.55 fatal 25.27 54.06 54.69 lean 1.662 2.452 2.470 * See Table # I Table # X Continued Daily production records 0. females #2-#12 for the fourth lactation, 5-18 days inclusive. Days ' 958 gfe 2&10 5111 g£12 Eean 5 .44 2.55 41.18 9.4 9.4 1.582 6 .67 1.54 1.58 1.875 7 .85 2.88 1.99 1.810 8 1.46 2.55 2.65 4 1.988 9 1.51 2.91 1.89 2.085 10 1.06 2.99 2.22 2.081 11 1.16 5.59 1.50 2.055 12 1.04 5.15 1.55 2.151 15 1.51 2.86 2.55 2.460 14 1.46 1.89 2.75 2.085 15 2.15 4.15 1.50 2.545 16 2.55 2.11 2.94 2.916 :17 2.19 4.05 2.42 2.555 18 2.08 4.06 5.45 5.088 165.1 19.69 40.62 29.65 lean 1.406 2.901 2.116 4 See Table # I -70- Table # XI lactation, 5-18 days inclusive. Daily production records of females #50-#57 for the first Iggy. 2#50 9#51 9#52 9&55 9#54 9#55 9 56 57 5 2.07 2.77 .61 2.49 .52 .55 .40 .55 6 2.84 2.56 1.90 2.98 .51 .68 .42 .21 7 2.55 5.72 1.70 2.65 .71 .89 .59 .42 8 1.45 5.52 1.20 5.20 .18 1.40 .57 .26 9 2.06 2.15 .70 2.49 1.42 1.52 .84 .51 10 1.51 2.94 .45 2.97 .21 2.51 .55 .58 11 .99 2.21 2.51 5.18 .51 .90 1.50 .09 12 .86 2.19 2.25 2.70 .29 2.51 1.05 .40 15 2.76 1.25 2.02 5.01 .24 2.06 .76 .82 14 5.47 2.57 1.54 2.65 1.55 .89 2.27 .85 15 2.11 .79 2.09 2.10 .72 1.7. 2.11 2.00 16 1.57 1.58 1.29 5.24 1.27 2.70 2.56 2.27 17 2.01 2.51 2.51 1.95 .59 1.15 5.04 1.72 18 2.21 2.58 2.62 2.64 .80 2.24 2.54 1.92 16541 27.82 52.52 25.17 58.21 9.52 21.58 18.95 12.16 ‘083 1e987 2e308 1e655 2.729 e665 1e521 1e353 e868 -71- Table # XII 'Daily production records of females #50 and #52 for the fourth lactation, 5-18 days inclusive. 221s gfég 9£52 5 1.25 i 1.19 6 .79 1.21 7 1.58 1.49 8 1.55 1.50 9 2.09 1.41 10 1.90 1.10 11 2.06 1.88 12 1.61 1.75 13 2.55 2.48 14 2.19 1.88 15 2.09 1.94 16 5.18 2.66 17 2.50 2.55 13 1.59 1.86 Total 26.51 24.56 lean 1.895 1.754 -72- Table # XIII Relationship between the amount ofmmilk consumed and the gain in weight ofyoung mice. Only litters of 7 young over a period of 14 days are included. Female Wt. of litter Total gain in Total amount Value (mother) at first wt. of young of milk of measurement. litter. consumed. K. (in grams) (in grams) (in grams) #2 19.97 6.04 29.68 .20 #5 18.76 16.95 42.55 .59 #4 18.51 10.59 54.77 .50 #5 17.65 7.29 15.45 .54 #6 19.56 9.00 54.80 .28 #8 17.51 7.55 25.88 .24 #9 17.61 20.81 51.51 .40 #10 17.55 9.77 57.91 .26 #11 19.11 15.17 45.90 .55 #12 17.18 15.01 57.21 .55 -75- Table # XIV Females vith.complete lactation records arranged in order of decreasing milk production on the basis of the total milk produced and their fourteen day samples. Complete Lactation , Fourteen Day Samples Female Lactation Total amount Female Lactation Total amount of milk of milk in grams. Vin grams. #11 2 91. 99 :11 2 45.90 #11 5 89.56 #11 5 45.60 #10 2 78.55 #10 2 57.91 #12 2 57.51 4 #12 2 57.21 #6 5 57.21 #4 2 54.77 #5 5 54.69 #6 5 28.55 #4 2 55.77 #5 5 27.66 #3 2 45.57 #8 2 25.88 #3 5 45.26 #8 5 15.78 #5 2 57.25 #5 2 15.45 -74- MICHIGAN STATE COI L '( .I‘IT'W'TV' v rv '.._. .V- vvvvfivyyv .717 VY' I 4 I! a . . 1 . . 4 . I V. _ 1 . 1 . _ _ . _ e I n .V . A . V _ 1 4 I a. V _ _ _ _ I I _ _ . I III . . , I 4 I .I D O _I...II> I II Ill: .6. . . . a I 1 I _ _ . 1 _ I V e I,. . 4 . . . V . ~ " a . 4 .I _ I I 4 a . 4 I I I I. .. I . _ _V . _ _ _ I _ . , . _ 1 a . l I I . . _ 1 1 I. . _ _ I .... . . . . v ... I . I .. 1 4 I _ . . _ V . .Ial...la 1 e I ll '9! n .I I I I In ll I _ . _ . , V; I I n. _ _ . I . . . e I _ V n _ V _ I. 1 _ k I a _ e I )I I I . O I . >g _ _ _ _ _ . _ C I . _ _ I In I 9 . . . II'I 7 III 9 1.\I.I r _ I'll . . V_ V V . V _ u . 4. \ I _ y _ ' _ q. p I . , I . I I . w . I V I I o e . 4 ' I . .I I 4 I II .4 I I .IO . a L _ . I . 0_ I V I0 . e o . 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Edd den ma ¢H NH 0H \m m n ma 5H ma ma Ha m b m nuuanaouq Pang. £593.03 vacuum on: 0303 no 8.35» hue 33.83 on» wnuunononqou guano noun—«pong aha 33334. enooom 23* 0305 «a «3%.: man 9.3.33 05 3333.3: 350 NH fi :53 cunts fit Ittn ‘m9 “I III)! -94- MICHIGAN STATE COLLEGE 1" ov .3553 an .3333 an» 3335 93 ammo .. .. J may 3.. .m‘ .mm .3 .mm .on .338 may. 13%....0 DEPARTM INT 0' HATHIIA? o . v o I. n.v . .V a . ... . .. cc. v V . I.. I .61-1 1. V.|,.V0 V. o 01 .4 . . _ . . ‘ n 04 ‘ a ..I _. O . .. .I.; II ...-I . l I} . , . . . I. 0‘ V I u IL I I . . 9.. _ n . I. 4 . v I I .o I v 9.. ._..w I I .h I .1 .I....Iili. I . 4 .di, . .Y‘l I . .- Op‘A b I I9? . I ...1 f .I 1.... V .II I .AL V L 041 o ._.. IJAV .- L I I I .++A I. _ I v o. A ‘ T I .O .1 1' I I I . .. I Y I>MVA U 0 .1 .V. . H 1 . I 9T1 I.§0 0A 7 OD , ._ I 9 0. I..v . I .. - _I. <4 41: . _.OIII $11 _ , _ I 0?} o. 9 I , vu « . + .1 I . OA“.QAA I I xlv AAA . I ._ . 9‘ AA 1 , 1' l .I ¥.> .I. . .« . oAvv . . 4. II I .o .- _v‘A o o . k . 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I.....46. m... .... ..J .F .IFI‘uIVAH .O A‘ c 1‘ I “Ital ..V“... . h I . ¢ 0%.: _OL.U.&I _ I h . .40.... e. ”P .4) I..... 1. $V>EFIIII -94- MICHIGAN STATE COLLEGE _ I . I nr I I ._ DI . III. I In. I J I I .. . 0‘ I I l JI I I, I "HI \ 4 . 4 . a I I I I. I _ I _ I_ _, .I. . l I . a , w- _ _ I . _ I . _ J I.I III]... lg II I I. I_ I I J .I _ I I" H . I .1. I I n . _ . I . _. ._I . I . I.I . . . I I I o I I .. I I I I. v I.._ . . . I I I I . _ __ . I III J. J IIIl_ In _ I I I IW . II I. II. _I I _ I I ...I ,.i I. I .....I . _ , I I .0 I . I I .v _‘ .I I. _. I . . I I. II I I . .- I .. 3 - . . .. . ,. I_I I .I« I a ‘ J . I I I . _ ,I_ . I I. I I I _ . _ _ ._ II I . .II _ . . I _ _ _. II _ I .n _ II I mm o ..I ...? . mI. _ u_ . . .. «I .. I. . . . I, . » O . ‘I I ... . f fig” I fictficflfimflhfi 0‘0 IO'nOHUflH “.99 :Huflfin II. II. 4 I n . . . . _.«.. _II.q II_< I n ,. M _I .I .I. I I. v I _ . _,. . . ”I n I . . II. I. ._ _ H I . I _I I . I. ., .I I. .I.I.I.II_. . I. I .II I I I. g . ._ II. . I, . I . I I - ._ .I , . ,0 I I. I‘ . I ._ u a _ ._ I . I o .. I I .-u I ._ .. n I. I I I v I _I A ...ol .I V II— _I , ,-H I_,I.I-I . I I I. QT . .5 I _ :_ ”a. w” . .Wt..-fim:wn..5. I . . . r I. I. m I __vI. II. - _ . I .I. w I I . - I. v v I. I . . n . _ II A H I ... , .I I.. I I I Anal. '. o . I I .,>.I,., H ..I. 3.. .. . . .v _I. _I. _‘I. any-IV. [.II II I T I ._LI . q ..I .I. . I. ., ._ I. I..I. .IJ.I . I.” ._JIIIIOEIIJY _I ' .II n. I _ _I . C .V . ..I I I. _I5_.. . — . .I “I. ’ w H u A I... I.I II_I I .0 I . I . . v .I I_ A ..I a r I I. _ .... I m. o I I I. _ V 0 q _ II I _ I v I u I r o O _ O I _ _ _ In . I. I I w“.- I . u I I I ”I .I. u- I v I..I.I I I )H‘I I I. “.IVI.M . I_ ._ I..I . . . -,I.v _ I . . _ .0. . a F I .o .F rt i I I) F).- lr * II, - IF? I b b) I 4 I A _.. I..; I .¢ DEPARTMENT OF HATHIHA‘ “UOUQ‘VO I. &. VI v I .I' I b .9 I v'I_ I I I I V I I . 7 ‘I v . I D ' .I I ._ . ' I I l. Ob v1lv I I D C v. o I v I I..- I .0 v . a. w. I. I v. I I I I I I .. I. _ T - II I v I v .II vI v fl. 0 V fI I .- _ I T. I v I r. _ I v 0 VI Y Q... II _ III _I v. I TI! 1 I . II. V. a . II I O .r. " I l v , O I I. I I I _ I 595- ~m om qu wu 3. «N om Emm aw . mH 3. NH 0H mm 5 o~&& o ’4 .wfiauwo ho 02.8ng 0193335 :3 3.230. a ‘ ¢ .. f ”K and .Mwa‘fiw 6* .nm .3.‘ sodium no #3433." go» 0.5 -530293 35.5 ‘ 3:338 UR. * 2H3.» -96- Part XVIII PLATE # ,I . The experimentsl cage is described on page 5. It¥s dimensions are 5% inches X8 inches X9 inches. 997- mm # 11 Rhotomicrograph ofva difoestrnus_smear of female #8 on June 27, 1954, stained with eosin and hematoxylin. X 560. 1. Strands of coagulated mucus. 2. Epithelial cells in various stages of degeneration. 3. Polymorphnuclear Ihc§§tes. :98- PLATE # III Photomicrograph of a pro-oestrous smear of females #8 on nJulyt 14. 1934, stained with eosin and hematoxylin. X 360. l. Lightly stained epithelial cells with small compact nuclei. :99- PLATE# _IV ‘Photomicrograph of an oestrous smear of female #8 on gJulyt 15, 1934, stained with eosin and hematoxylin. X 360. l. Cornified, eosinophil, non-nucleated epithelial cells. -100- ;: ‘3 . '56‘ . f!» "A: PLATE #‘V Photomicrograph of'apmetafoestrous smear of female #8 on fiJulyt 17,_1934, stained with eosin and hematoxylin. X 360. _ 1. Typical nucleated epithelial cell. 2. Mass of mucus. 3. Polymorphonuclear leucoytes. {101- ,PLATE # VI ,Photomicrograph of an oestrous smear of female #2 on LJuIyt 13,.1934, stained with eosin and hematoxylin. X 560. l. Cornified, eosinophil, non-nucleated epithelial cells. 2. products of cell decomposition. 1102- PLATE # VII fhotomicrograph of an oestrous smear of female #3 on .Julyi 20,'1934, stained with eosin and hematoxylin. X 360. 1. Cornified, eosinophil, non-nucleated epithelial cells. '— *VU— PLATE # VIII Photomicrograph of an oestrous smear of female #5 on my: 16, .1934, stained with eosin and hematoxylin. x 360. l. Cornified, eosinophil, non-nucleated epithelial cells. _.PLATE # Ix 2hotomicrograph of an oestrous smear % of female #9 on gxulyV 16, 1954, stained 1 with eosin and hematoxylin. x 360. l. Cornified, eosinophil, non-nucleated . epithelial cell. . . . 2. Typical nucleated epithelial cell indicating the later stages of oestrus. 3. Products of cell degeneration. 2105- PLATE # X fhotomicrograph of an oestrous smear_ of» females #10 on July 20, .1934, stained ‘with eosin and hematoxylin. X 360. -l. Cornified, eosinophil, non-nucleated epithelial cells. 2. Products of cell degeneration. ROOM USE ONLY .Y2033 ”/63 SE} a ~ 4 a . 1 n ‘ I ‘ u . 0 . n , . . . 1 . ’ I v . o . ‘ . . . . K n . . . v . - . J . . . . . l u v _ ‘ . , .. , . - y l . , _ ‘ 1 . . x u k . _. . . - ‘ . 1‘ I a .t l ‘.. . ' . I ‘ . .. ‘ I . . A . . \ . . u - ‘ o . . ‘ i 1 . 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