Behavior Genetics, Vol. 6, No.

4, 1976

Analysis of Truncated Distributions: Mating Speed in Drosophila melanogaster
Maurice A. Dow

Received 24 Aug. 1975-Fina/17 Nov. 1975

The distribution of mating speeds in wild-type Drosophila melanogaster is shown to be log normal. The analysis of mating speeds by methods for trun cated distributions is validated, and unbiased estimates of the mean mating speed, the variance of mating speed, and the proportion of flies capable of ever mating are produced. In pair matings, not all pairs are capable of copulating, given even a 7-day mating period.
KEY WORDS: Drosophila meianogaster; data analysis; mating speed; truncated distributions.

INTRODUCTION Many of the phenotypes of interest to behavior geneticists have truncated distributions (e.g., ejaculation latencies in mice, mating speeds in Drosophila-not all animals react within a reasonable time limit and it is not certain that all animals would react given enough time). This causes considerable difficulties in data analysis and results in each investigator preferring his own, possibly statistically invalid method of analysis (Fulker, 1966; Manning, 1961; Parsons, 1964; Spiess and Langer, 1964; Spiess et al., 1966). The bias involved in comparisons of uncorrected means of truncated distributions has been pointed out by Nagylaki (1973). Olshen (1970) reviewed the methods used for the analysis of Drosophila mating speeds and concluded that the probit plot vs. time in logarithms (Manning, 1961; Spiess and Langer, 1964) was the most suitable, on the assumption that eventually all animals would copulate. However, Bliss
Supported by the National Research Council of Canada.

Department of Zoology, University Edinburgh, Edinburgh, Scotland.


Behavior Genetics, Vol. 6, No.4, 1976


1976 Plenum Publishing Corporation, 227 West 17th Street, New York, N.Y. 10011. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or l:>y any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission of the publisher.



(1967) has described a statistical method for analyzing truncated distribu tions that allows this assumption to be tested and that estimates the unbiased mean mating speed, the unbiased variance of mating speed, and the number of animals capable of ever copulating in a given sample. This method requires as input the sample mean, the sample second moment (the variance with divisor N, not N - l), and the limiting value (time limit) of the observed mating speeds. It also requires that the mating speeds be measured in units that are normally distributed. As a first approximation, the data can be checked for normality using probit plots (Wright, 1968) or computer programs such as A3.1 (Sokal and Rohlf, 1969) and transformed appropriately. The adequacy of any transformation needed can be tested using the results produced by the analysis. The procedure is described in detail in Bliss (1967, p. 162) and a worked example is given in the Appendix. The appropriateness of using the method with logarithmically transformed mating speeds in wild-type Drosophila melanogaster is tested. This method has been used by von Schilcher and Manning (1975), although they did not discuss its validity.

In the first experiment, 135 single pairs of 4-day-old outbred virgin wildtype flies were introduced, without etherization, into shell vials (20 by 76 mm) containing 7 ml of propionic acid medium and observed for 40 min each. The results of this experiment were used to test the assumption that mating speeds are lognormally distributed. The distribution of mating times predicted from the estimates of the "true" mean, variance, and number of pairs ever copulating obtained from the analysis for truncated data was compared to the observed distribution by a 2 goodness-of-fit test. The procedure for the second experiment followed that of the first experiment except that all pairs were observed for 1 hr and the vials were kept for a further 7 days so that an estimate of the total number of pairs ever mating could be made. This estimate was corrected for sterility based on the percentage sterility found in the pairs that were seen to mate during the 1-hr observation period. A number of replicates of various sizes were run. This experiment was used to compare the observed number of pairs capable of ever mating to the number predicted by the analysis for trun cated data.

Table I reveals that the logarithmic transformation gives a satisfactory fit to the observed mating speeds, on the assumption that 123.5 of the 135

Mating Speed in D. melanogaster 387

Table I. Frequency Distribution of Log Mating Speeds" Mating speed interval (log min) - oo-0.59 0.60-0.73 0.74-0.83 0.84-0.92 0.93-1.00 1.01-1.08 1.09-1.17 1.18-1.27 1.28-1.41 1.42- 00 "x2 = Frequency Observed 12 II 14 12 15 14 13 10 6 16 Expected 12.16 12.25 12.40 12.26 12.67 12.67 12.26 12.40 12.25 12.16

6.16, df = N - 3 = 7, P > 0.50, predicted mean
1.00 log min, predicted standard deviation = 0.32 log min, predicted number of pairs ever mating = 123.5.

pairs would ever mate. This is only 3.5 pairs more than were observed to mate within 40 min. As shown in Table II, the number of pairs capable of ever copulating is adequately predicted. On the average, only 89% of the pairs ever mated.

The application of the analytical method for truncated distributions to wild-type D. melanogaster mating speeds has indicated that mating speed is lognormally distributed and that not all pairs are capable of mating. As this method is simple and well documented (Bliss, 1967), it should prove useful
Table 11. Comparison of the Observed and Predicted Number of Pairs Capable of Ever Copulating Total number of copulations Sample size 50 30 32 126 50 288 Observed 4 25 5 2 1 120 4 6 2 5 Predicted 42. 30 7 20. 2 119. 544. 4 256. 8

388 Dow

in future work on Drosophila mating speeds. It is also applicable in the analysis of other behavioral phenotypes (e.g., ejaculation latencies in mice) once specifically validated for each case. APPENDIX

In a sample of 120 pairs, 27 (N) mated within 60 min. The mating speeds (ms), originally measured in minutes, were transformed to logarithms for the following calculations.

Total of mating speeds (2:ms) = 38.0839 Total of squared mating speeds (2:ms2 Observedmean(m 1
= 55.3005

= 55.3005 - (1450.3834/27) = 1.5826 Second moment (m2) = ssj N = 1.5826/27 = 0.0586 Time limit (Yo) = 60 min transformed to logarithm = 1.7782

) = 2:msjN = 38.0839/27 = 2 1.4105 Sum (2:ms)2/ N of squares (ss) = 2:ms -



m2j(m1- Y0)2 = 0.0586/(1.4105 -



Use H to look up {J in Table A.11 (Bliss, 1967)
{j = 0.3575

Estimated mean (m) = m1 - {J (m1 - Yo) = 1.4105 - 0.3575(1.4105 - 1.7782) = 1.5419 2 Estimated variance (.P) = m2 + {j (m1 - Y ) 0 2 = 0.0586 + 0.3575(1.4105- 1.7782) = 0.1069 Estimated standard deviation (s) = .y'¥ =0.3270 For mating speeds, the distribution is truncated at the upper (right-hand) side; therefore Xo = (m - Yo)/S = (1.5419- 1.7782)/0.327 = -0.7224 Use Xo to look up v 1

in Table A.13 (Bliss, 1967)
v1 = 5.3347

388 Dow

Varianceofthemean(Vm) = v1(s2)/N = 0.0211

= 5.3347(0.1069)/27

Mating Speed in D. melanogaster


Standard error of the mean (se) UseX0 to

= =



look up fo in Table A.4 (Bliss, 1967)
Po = 0.236

Estimated number of pairs to ever mate (IV)= Nj(l -Po) = 35.3 Final Results Estimated mean mating speed = 1.5419 log min Estimated standard deviation = 0.327 log min Estimated standard error of the mean = 0.1454 log min Estimated number of pairs to ever mate = 35.3 REFERENCES
Bliss, C. I. (1967). Statistics in Biology, Vol. I, McGraw-Hill, New York, p. 162. Fulker, D. W. (1966). Mating speeds in male Drosophila melanogaster: A psychogenetic analysis. Science 153:203-205. Manning, A. (1961). The effects of artificial selection for mating speed in Drosophila me/ana Nagylaki, T. (1973). Sampling truncated distributions. Behav. Genet. 3:193-196. Olshen, V. A. S. (1970). Some methods of analysis for data on mating latency. Ph.D. thesis, Yale University. Parsons, P. A. (1964). A diallel cross for mating speeds in Drosophila melanogaster. Genetica Sokal, R. R., and Rohlf, F. J. (1969). Biometry, Freeman, San Francisco. Spiess, E. B., and Langer, B. (1964). Mating speed control by gene arrangement carriers in Spiess, E. B., Langer, B., and Spiess, L. D. (1966). Mating control by gene arrangements in von Schilcher, F., and Manning, A. (1975). Courtship song and mating speed in hybrids between Drosophila melanogaster and Drosophila simulans. Behav. Genet. 5:395-404. Wright, S. (1968). Evolution and the Genetics of Populations, Vol. I, University of Chicago Press, Chicago.
Drosophila pseudoobscura. Genetics 54:1139-1149. Drosophila persimilis. Evolution 18:430-444. 35:141-151. gaster. Anim. Behav. 9:82-92.

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