some data were not expected to be normally distributed, e.g. durations of behaviour Ž
. patterns are limited to 300 s 5 min , favouring nonparametric statistics. Therefore, all
data are presented as medians instead of means. Total duration and frequencies of post-meal behaviour were tested for outliers using the ‘Explore: Statistics’ function of
SPSS.
Sequences of events of 5 min post-meal behaviour of MFF or LFF were obtained by generating transition matrices for each group of cats. Matrices were edited and tested for
Ž significant transitions in Matman version 1.0 Noldus Information Technology, Wa-
. Ž
. geningen, the Netherlands . Significant transitions
p F 0.05–0.001 were detected
Ž .
using the chi-square x 2 test followed by the adjusted residuals procedure in Matman. Ž
. Sign tests Ferguson, 1981, pp. 401–402 were used to test for differences between
Ž .
the amounts of MFF and LFF, and normal food before or after MFF NF-MFF and Ž
. normal food before or after LFF NF-LFF eaten per cat within the three groups. Per
group, it was scored in how many cases a cat ate more of MFF than of LFF and more of NF-MFF than of NF-LFF. Between groups x 2 tests were used.
3. Results
3.1. General Ž
. To test for order effects
MFF before LFF and vice versa total duration and
frequencies of post-meal behavioural patterns for each of these subgroups for each of the four food items were tested against one another. In groups B and C, no significant
Ž .
differences were found p 0.05; Mann–Whitney U-test . Group A contained too few
cats to run a meaningful analysis, but no differences appeared to be present upon visual inspection of the data. Therefore, results of the cats within one group were pooled.
Furthermore, total duration and frequencies of post-meal behaviour were tested for outliers. Special attention was paid to the four cats that did not comply with all selection
criteria. No outliers were detected. 3.2. Amount of the food items eaten and latencies to eating MFF and LFF
Ž In group A, more MFF than LFF tended to be eaten per cat sign-test: z s 1.5,
. p F 0.14 whereas no difference was observed between the amount of NF-MFF and
Ž .
NF-LFF eaten per cat z s 0, n.s.; Table 3 . In group B, no differences were detected Ž
. between the amount of MFF and LFF eaten per cat
z s 1.15, n.s. , or NF-MFF and Ž
. NF-LFF eaten per cat z s 0.58, n.s. . In group C, more MFF than LFF was eaten per
Ž .
Ž .
cat z s 2.67, p F 0.01 as well as more NF-LFF than NF-MFF z s 2.04, p F 0.05 . No differences were observed as to the amounts of MFF eaten per cat between
Ž .
groups: group A versus group B x
2 s 0.66, df s 2, n.s. , group B versus group C Ž
. x
2 s 1.68, df s 2, n.s. . In contrast cats in group B ate significantly more of LFF than Ž
. cats in group C x 2 s 9.49, df s 2, p F 0.01 . Although cats in group B ate also more
Ž of LFF than cats in group A this difference did not reach significance
x 2 s 3.18,
. df s 2, n.s. . Although cats in group B tended to eat more NF-MFF and NF-LFF than
Table 3 Number of cats eating all, a part or nothing of the four food items during testing expressed as percentages of
total number of cats per group Group
No. of cats in group MFF
LFF All
Part Nothing
All Part
Nothing A
5 100.0
0.0 0.0
20.0 20.0
60.0 B
8 87.5
12.5 0.0
62.5 0.0
37.5 C
10 60.0
40.0 0.0
0.0 30.0
70.0 NF-MFF
NF-LFF All
Part Nothing
All Part
Nothing A
5 60.0
40.0 0.0
40.0 60.0
0.0 B
8 62.5
25.0 12.5
75.0 25.0
0.0 C
10 20.0
40.0 40.0
40.0 60.0
0.0
Ž cats in group C this did not reach significance x 2 s 3.58, df s 2, n.s. and x 2 s 2.22,
. df s 2, n.s. .
In group A, it was not possible to measure the latencies for eating MFF or LFF precisely, since these food items were presented as the second food item, and therefore
the exact moment the cat was presented the food item was never the same. In group B, Ž
. Ž
the latency to eating MFF median: 0.0 s was shorter than to eating LFF 170.1 s; .
Wilcoxon matched-pairs signed ranks test: z s y2.52, p F 0.05 . The same was true Ž
. for group C: MFF, 2.2 s versus LFF, 300.0 s
z s y2.81, p F 0.01 . There were no Ž
differences in latency to eating either MFF or LFF between groups B and C Mann– .
Whitney U-tests, n.s. . Of the five cats of group B that did eat LFF, 81.6 of the time spent before eating
Ž .
LFF median value consisted of sniffing and licking at the food. The three cats of group C that ate from the LFF were spending 84.7 of the latency period by sniffing and
Ž .
licking at it median value . The percentages of time spent on licking or sniffing at LFF Ž
. of group B and C were not significantly different Mann–Whitney U-test, n.s. .
3.3. Sequences of behaÕioural patterns Post-meal behavioural patterns were recorded as a sequence of events. Transition
probabilities of behavioural patterns for each group of cats were calculated for both MFF and LFF. There was a strong resemblance between the resulting sequences for the
different groups in terms of general patterns, and the behavioural events at the beginning and end of the sequences. Therefore, the results of all the cats were pooled resulting in
one sequence for post-meal behaviour of MFF and one sequence for post-meal be-
Ž .
Ž .
haviour of LFF. In the matrix of MFF in total 441 cells , 200 cells 45.4 had a value Ž
. Ž
of zero and 147 cells 33.3 had a value of F 5. In the matrix of LFF in total 441 .
Ž .
Ž .
cells , 133 cells 30.2 had a value of zero and 193 cells 43.8 had a value of F 5.
To test if the resulting sequences were however representative for common cat post-meal behaviour, ‘split half tests’ were performed. This means that transition probabilities were
Ž calculated again, using the results of only half the number of animals
n s 11 and .
n s 12 cats; the distribution of cats over the two halves was randomly assigned , leading to two ‘new’ sequences for both MFF and LFF. The resulting split-half sequences
resembled the original sequences. Therefore, it can be assumed that if a larger group of cats was used and thus a higher number of behavioural events would emerge, the
sequence would not have changed very much either.
Ž The sequences of MFF and LFF are shown in Fig. 1 both matrices were significant:
x 2 s 2064.4, df s 341, z s 38.2, p F 0.001 and x 2 s 1692.5, df s 419, z s 29.3,
. p F 0.001, respectively, Ferguson, 1981, p. 204 . Both sequences started with activities
Ž around the feeding bowl such as lick r sniff food 56.5, only displayed when LFF was
. Ž
presented , lick r sniff feeding bowl 26 when presented LFF and 47.8 when
. Ž
presented MFF and sniff floor 21.7 when presented MFF; this pattern could be displayed anywhere in the room but was most frequently seen in close proximity of the
feeding bowl; percentages were calculated as the number of times the patterns were marked as the first event in the individual sequences as a proportion of the total number
Ž ..
of individual sequences n s 23 . The activities around the feeding bowl were followed by a period of walking around, exploration of the room and interaction with the owner
andror the observer. Near the end of the sequences a period of grooming occurred, which was normally done while the cat sat, therefore sit up is closely associated with
the different grooming behaviours. The sequence was usually ended by sit or lie down Ž
. Ž
17.4 and 21.7 when presented MFF or LFF, respectively , sit up 30.4 when .
Ž .
presented MFF , lick paw 8.7 when presented MFF or groom chest r groom body Ž8.7 when presented LFF; percentages were calculated as the number of times the
patterns were marked as the last event in the individual sequences as a proportion of the Ž
.. total number of individual sequences n s 23 .
The MFF and LFF sequences showed a number of differences. Lick nose was only present in the LFF sequence. Furthermore, the LFF sequence started with either
lick r sniff feeding bowl or lick r sniff food while the MFF sequence started with lick r sniff feeding bowl only. Considering the fact that only 10 of the 23 cats ate LFF, it
Ž .
was decided to see whether the behavioural sequence of the cats n s 10 that ate LFF Ž
. all of it or only a part
was different from the behavioural sequence of the cats Ž
. n s 13 that did not eat any LFF in terms of, e.g. these initial behavioural patterns.
Ž Both matrices were significant: x 2 s 986, df s 419, z s 15.5, p F 0.001 cats eating
. Ž
LFF, n s 10 and x 2 s 1045.9, df s 419, z s 16.8, p F 0.001 cats not eating LFF, .
Ž .
n s 13 . The sequence of the cats that ate no LFF at all LFF refusal sequence started Ž
with lick r sniff food 100; lick r sniff feeding bowl was only recorded twice in these .
Ž .
cats , while the sequence of the cats that did eat LFF LFF consumption sequence Ž
started with lick r sniff feeding bowl 60; lick r sniff food was only recorded three .
Ž .
times in these cats or sniff floor 20 similar to after eating MFF. Furthermore, the ‘LFF consumption sequence’ but not the ‘LFF refusal sequence’ resembled the ‘MFF
Ž .
consumption sequence’ for example in grooming behaviour not shown . It should be noted that lick nose was present in both LFF sequences: it occurred five times in cats
that ate LFF, and 25 times in cats that did not.
3.4. Total duration and frequencies of behaÕioural patterns In Table 4, total duration and frequencies of behavioural patterns for the different
food items are presented per group. For ease of presentation and subsequent discussion, Table 4A contains only those behavioural patterns that showed consistent differences
over the groups and food items, i.e. the patterns that were in line with our hypothesis. The data of the remaining behavioural patterns are listed in Table 4B. Neither the data of
MFF and NF-LFF, nor the data of LFF and NF-MFF were compared with one another since this would not give any extra useful information. The most relevant differences
between food items within the groups will be summarised per group.
3.4.1. Group A The cats of group A, which were normally fed one to three meals a day and which
started both observations with their normal food, showed no differences in behavioural patterns between NF-MFF and NF-LFF. In group A, total duration and frequency of
groom face were significantly increased after MFF when compared to after LFF, and to after NF-MFF. Except for the frequencies of MFF and LFF the same was true for
lick r sniff feeding bowl. The frequency of groom chest r groom body was significantly increased after MFF when compared to after NF-MFF.
3.4.2. Group B In group B, the cats were normally fed one to three meals a day and started the
observations with one of the experimental food items. Total duration of lick r sniff feeding bowl was significantly increased after MFF compared to after LFF. The
frequency of lick r sniff feeding bowl after MFF was significantly higher compared to after NF-MFF. The frequency of sniff floor was significantly increased after MFF when
compared to after NF-MFF as well as after LFF when compared to after NF-LFF. Total duration of sniff floor after LFF was significantly higher than after MFF. Finally, total
duration of groom chest r groom body after NF-MFF was significantly increased when compared to after MFF.
3.4.3. Group C Group C consisted of cats fed ad libitum by their owners and started the observations
with one of the experimental food items. Total duration and frequency of lick r sniff food after LFF were significantly increased when compared to after MFF, and to after
Fig. 1. Post-meal transition probabilities between behavioural patterns after different food items. Abbreviations of patterns are found in Table 2. Frequencies are shown between brackets. Sequences start at the top
Ž .
Ž .
underlined behavioural patterns and end at the bottom underlined behavioural patterns . Panel A: Presumed Ž
. sequence of patterns after MFF Sheba trout . Data of groups A, B and C combined to reveal one sequence
Ž .
Ž ns 23 cats, 1661 transitions, see text . Panel B: Presumed sequence of patterns after LFF common brand
. Ž
with drops of orange essence . Data of groups A, B, and C combined to reveal one sequence ns 23 cats, .
1629 transitions, see text . P F 0.001;
P F 0.01; P F 0.02;
P F 0.05.
NF-LFF. Total duration of lick r sniff feeding bowl was significantly increased after MFF and NF-LFF compared to after NF-MFF. The frequency of sniff floor was
significantly increased after MFF when compared to after NF-MFF. Total duration of sniff floor after NF-MFF as well as after LFF and frequency of sniff floor after LFF
were significantly increased when compared to after NF-LFF. Total duration and frequency of lick nose after LFF were significantly increased with respect to after MFF
and to after NF-LFF. There was a significant increase in total duration and frequency of lip lick after MFF when compared to after LFF and in frequency after MFF when
compared to after NF-MFF. Total duration and frequency of lip lick after NF-LFF were significantly increased when compared to after LFF. The frequency of lip lick was also
significantly higher after NF-LFF than after NF-MFF. Total duration and frequency of
Table 4 Median values of total duration and frequencies of behavioural patterns per food item per group
Ž . A
Total duration Frequencies
MFF LFF
NF-MFF NF-LFF
MFF LFF
NF-MFF NF-LFF
Group A Lf
0.0 17.7
0.0 0.0
0.0 2.0
0.0 0.0
a a
Lb 15.2
0.0 0.0
0.0 4.0
0.0 0.0
0.0 Ln
0.0 1.6B
0.0 0.0
0.0 1.0B
0.0 0.0
Ll 30.5
14.3 27.4
32.2 12.0
10.0 13.0
14.0
a a
Gf 63.7B
10.8 6.0
16.9 5.0BB
2.0 1.0
3.0 Group B
Lf 0.0
0.0 0.0
0.0 0.0
0.0 0.0
0.0
a
Lb 39.3
9.3 3.5
5.9 3.5
1.0 1.0
1.0 Ln
0.0 0.0
0.0 0.0
0.0 0.0
0.0 0.0
Ll 46.5
32.3 31.4
39.0 14.5
13.5 12.0
15.0 Gf
15.2 5.9
42.2 22.1
1.0 1.5
2.5 2.0
Group C
† †
Lf 0.0
19.7 B
0.0 0.0
0.0 2.5
0.0 0.0
a ‡
Lb 9.7
0.6 0.0
2.2 0.5
0.5 0.0
0.5
† †
Ln 0.0
2.3 0.0
0.0 0.0
1.0 0.0
0.0
† a
†,‡
Ll 47.1
10.3 29.7
36.7 16.5
5.0 11.0
14.5 Gf
31.2 0.0
13.3 29.8
2.5 0.0
2.0 2.0
Ž . B
Total duration Frequencies
MFF LFF
NF-MFF NF-LFF
MFF LFF
NF-MFF NF-LFF
Group A Sf
11.6 2.6
2.8 2.7
4.0 1.0
1.0 1.0
Wd 4.4
5.3 7.5
5.6 2.0
2.0 5.0
2.0 Wu
0.0 0.0
9.1B 0.0
0.0 0.0
3.0B 0.0
Jo 1.3
2.7 2.4
0.0 1.0
1.0 1.0
0.0
a
St 5.0
7.8 18.3 B
8.7 4.0
4.0 11.0B
3.0 Ft
0.0 0.0
1.7 0.0
0.0 0.0
1.0 0.0
Other 6.7
9.3 18.8
3.2 1.0
3.0 9.0
1.0 Rub
0.0 0.0
7.7 0.0
0.0 0.0
3.0 0.0
So 0.0
2.7 11.1
4.6 0.0
1.0 3.0
1.0
a
Lm 0.9
4.9 3.2
0.0 1.0
2.0 2.0
0.0 La
7.2 11.1
20.1 10.1
3.0 4.0
6.0 3.0
Lo 0.0
6.5 2.7
2.5 0.0
3.0 1.0B
1.0 Su
15.3 22.1
32.4 48.1
11.0 10.0
8.0 9.0
Lp 9.8B
17.8 7.8
16.0 2.0B
2.0 2.0
4.0
a
GcrGb 15.4
3.0 10.4
18.9 5.0BB
1.0 3.0
2.0 SirLd
0.0 40.8
0.0 3.6
0.0 4.0
0.0 1.0
Group B
a †
Sf 6.5
7.8 0.0
0.0 3.5
2.0 0.0
0.0 Wd
10.0 15.0A
4.7 12.5
4.5 4.0
1.0 3.0
†,‡ a
a
Wu 4.4
5.6 0.0
1.6 2.0
3.0 0.0
0.5
Ž .
Table 4 continued Ž .
B Total duration
Frequencies MFF
LFF NF-MFF
NF-LFF MFF
LFF NF-MFF
NF-LFF Group B
a
Jo 2.6A
1.6 0.8
1.5 1.5
1.0 0.5
1.0
a
St 14.8
13.2 3.2
9.8 6.5
5.5 1.5
3.5
a
Ft 1.8
0.0 0.0
0.0 1.0
0.0 0.0
0.0
a †
†
Other 11.2
26.2 2.9
4.0 3.0
7.0 1.0
2.0 Rub
0.0 0.0
0.0 0.0
0.0 0.0
0.0 0.0
a a
So 10.0
7.0 0.0
0.0 2.5
2.5 0.0
0.0 Lm
2.6 4.5
0.0 1.0
1.5 2.5
0.0 0.5
a
La 7.2
11.5 11.9
19.4 2.5
4.5 3.0
5.5
†
Lo 2.2
5.3 0.0
0.0 1.0
2.0 0.0
0.0 Su
25.3 17.2
27.1 52.5
6.0 8.0
6.0 11.5
Lp 0.0
2.0 3.4
7.1 0.0
1.0 1.0
1.5
a
GcrGb 9.1
0.0 23.5
11.2 2.0
0.0 2.5
2.5 SirLd
3.6 0.0
3.0 0.0
1.0 0.0
1.0 0.0
Group C
†,‡ a
†
Sf 4.0
4.8 1.7
0.0 2.0
2.0 0.5
0.0
†
Wd 6.7
16.3 6.5
6.1 2.0
5.0 3.0
2.0
† †
Wu 8.1
15.4 10.3B
5.8 2.5
4.5 3.0B
2.0 Jo
3.7 4.1
4.0 1.8
1.5 2.0
1.0 1.0
† †
St 7.7
20.1 11.7
4.5 3.0
6.5 4.5
2.5
†,‡
Ft 1.0
3.0 1.6
0.0 0.5
1.5 1.0
0.0 Other
10.3 9.8
11.0B 12.1
2.5 3.0
2.5 4.5
† †
Rub 0.0
2.6 0.0
0.0 0.0
1.0 0.0
0.0
†
So 9.6
9.7 7.1
1.4 3.5
3.5 1.5
1.0
a
Lm 2.3
3.6 1.4
6.4B 1.5
2.0 1.0
1.5 La
8.5 21.3
14.9 9.6
2.0 4.0
3.0 3.0
‡
Lo 0.0
3.5 0.0
4.5 0.0
1.0 0.0
2.0 Su
11.3 35.6
18.2 37.8
4.0 7.5
5.5 8.5
†
Lp 4.8
0.0 0.0
5.6 1.0
0.0 0.0
2.0 GcrGb
12.2 2.0
7.8 10.7
2.0 1.0
1.5 1.5
SirLd 0.0
0.0 7.1
0.0 0.0
0.0 0.5
0.0 Ž
. Symbols indicate the level of significance between food items Wilcoxon matched-pairs signed-ranks test .
Ž .
Capitals refer to differences between food items of groups Mann–Whitney U-test ; single capital: pF 0.05, Ž .
double capital: pF 0.01. Capitals are placed near the highest value. In A , patterns are shown of which the Ž .
changes over food items and groups are consistent with the hypothesis formulated in Section 1, in B patterns Ž
. which are not see text for discussion . For abbreviations of behavioural patterns, see Table 2.
Ž .
LFF MFF pF0.05 . Ž
. LFF MFF pF0.01 .
a
Ž .
NF-MFF MFF pF0.05 .
†
Ž .
NF-LFF LFF pF0.05 .
‡
Ž .
NF-LFF NF-MFF pF0.05 .
groom face were significantly increased after MFF when compared to after LFF. Finally, after NF-LFF, the frequency of lick paw was significantly higher than after
LFF.
3.5. Differences between the groups The frequencies and total duration of lick paw and groom face and the frequency of
groom chest r groom body after MFF were higher in group A compared to group B. Furthermore, total duration and frequency of lick nose were increased after LFF in
group A when compared to group B. In group C, total duration of lick r sniff food after LFF was increased when compared to group B.
4. Discussion
