Behavior Genetics, Vol. 6, No.

4, 1976

Analysis of Truncated Distributions: Mating Speed in

Drosophila melanogaster
M a u r i c e A. D o w x
Received 24 Aug. 1975--Final 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 truncated 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
M a n y 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.
1 Department of Zoology, University of Edinburgh, Edinburgh, Scotland.
385
9 1976 Plenum Publishing Corporation, 227 West 17th Street, New York, N.Y. 10011. No part of
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(1967) has described a statistical method for analyzing truncated distributions 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 - 1), 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 yon
Schilcher and Manning (1975), although they did not discuss its validity.
MATERIALS AND M E T H O D S
In the first experiment, 135 single pairs of 4-day-01d outbred virgin
wild-type flies were introduced, without etherization, into shell vials (20 by
76 ram) containing 7 ml of propionic acid medium and observed for 40 rain
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 X~ 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 truncated data.
RESULTS
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

Table

1.

387

Frequency

Distribution
Speeds ~

of

Log

Mating

Mating speed
interval
(log min)

Frequency
Observed

Expected

- ~-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- ~

12
11
14
12
15
14
13
10
6
16

12.16
12.25
12.40
12.26
12.67
12.67
12.26
12.40
12.25
12.16

a X2 = 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.
DISCUSSION

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 II. Comparison of the Observed and Predicted
N u m b e r of Pairs Capable of Ever Copulating

Total number
of copulations
Sample
size

Observed

Predicted

50
30
32
126
50

45
25
21
120
46

42.7
30
20.2
119.5
44.4

288

257

256.8

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in future w o r k on D r o s o p h i l a m a t i n g speeds. It is also a p p l i c a b l e in the

analysis of other b e h a v i o r a l p h e n o t y p e s (e.g., e j a c u l a t i o n latencies in mice)
once specifically v a l i d a t e d for each case.

APPENDIX
Example
In a s a m p l e of 120 pairs, 27 (N) m a t e d within 60 rain. T h e m a t i n g
speeds (ms), originally m e a s u r e d in minutes, were t r a n s f o r m e d to
l o g a r i t h m s for the following calculations.
T o t a l of m a t i n g speeds ( ~ m s ) = 38.0839
T o t a l of s q u a r e d m a t i n g speeds ( ~ m s 2) = 55.3005
O b s e r v e d m e a n ( m l ) = Z m s / N = 38.0839/27 = 1.4105
S u m of squares (ss) = Z m s 2 - ( Z m s ) 2 / N
= 55.3005 - (1450.3834/27) = 1.5826
S e c o n d m o m e n t (m2) = s s / N = 1.5826/27 = 0.0586
T i m e limit (I7o) = 60 rain
t r a n s f o r m e d to l o g a r i t h m = 1.7782
H = m 2 / ( m l - Yo) 2 = 0.0586/(1.4105 - 1.7782) z
= 0.4337

U s e H to l o o k up 0 in T a b l e A. 11 (Bliss, 1967)
0 = 0.3575
E s t i m a t e d m e a n (rh) = m l - 0 (m~ - Yo)
= 1.4105 - 0 . 3 5 7 5 ( 1 . 4 1 0 5 - 1.7782)
= 1.5419
E s t i m a t e d v a r i a n c e (52) = m2 + 0 ( m l - Yo)~
= 0.0586 + 0.3575(1.4105 - 1.7782) 2
= 0.1069
E s t i m a t e d s t a n d a r d d e v i a t i o n (5) =
= 0.3270
F o r m a t i n g speeds, the d i s t r i b u t i o n is t r u n c a t e d at the u p p e r ( r i g h t - h a n d )
side; t h e r e f o r e
X o --- (rh - Yo)/~ = (1.5419 - 1.7782)/0.327
= -0.7224

U s e Xo to l o o k up v~ in T a b l e A.13 (Bliss, 1967)

vl = 5.3347
V a r i a n c e of the m e a n (Vm) = v~(~2)/N = 5.3347(0.1069)/27
= 0.0211

Mating Speed in D. melanogaster

389

S t a n d a r d error of the m e a n (se) =

= 0.1454
U s e Xo to look up Po in T a b l e A.4 (Bliss, 1967)

Po = 0.236
Estimated n u m b e r of pairs to ever m a t e (N) = N / ( 1 - Po)
= 35.3

Final Results
Estimated
Estimated
Estimated
Estimated

m e a n m a t i n g speed = 1.5419 log m i n

s t a n d a r d deviation = 0.327 log m i n
s t a n d a r d error of the m e a n = 0.1454 log m i n
n u m b e r of pairs to ever m a t e = 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 rnelanogaster: A psychogenetic
analysis. Science 153:203-205.
Manning, A. (1961). The effects of artificial selection for mating speed in Drosophila melanogaster. Anirn. Behav. 9:82-92.
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
35:141-151.
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
Drosophila persimilis. Evolution 18:430-444.
Spiess, E. B., Langer, B., and Spiess, L. D. (1966). Mating control by gene arrangements in
Drosophila pseudoobscura. Genetics 54:1139 1149.
yon 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 oJ Populations, Vol. I, University of Chicago
Press, Chicago.