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Behavioural Brain Research 267 (2014) 119–125

Contents lists available at ScienceDirect

Behavioural Brain Research


journal homepage: www.elsevier.com/locate/bbr

Research report

Loss of motor coordination in an aging mouse model


Erika N. Allen, Jane E. Cavanaugh ∗
Division of Pharmaceutical Sciences, Mylan School of Pharmacy, Duquesne University, 600 Forbes Avenue, Pittsburgh, PA 15282, United States

h i g h l i g h t s

• Aged C57Bl/6 mice exhibit a loss of motor performance.


• Weight and food intake are not confounding factors for behavioral tests.
• Middle-aged mice exhibit a loss of motor coordination between 11 and 12 months.

a r t i c l e i n f o a b s t r a c t

Article history: With age, there is an increase in motor deficits that leads to an increased incidence of slips and falls.
Received 20 November 2013 As the elderly population continues to grow, there is a need for aging models and research that focus
Received in revised form 12 March 2014 on behavioral deficits that occur with normal, non-diseased aging. The present study was designed to
Accepted 17 March 2014
examine the appropriateness of C57Bl/6 male mice as aging animal models using the challenging beam
Available online 25 March 2014
and cylinder tests to measure motor coordination and spontaneous activity, respectively. Using young
(2–4 mo), middle-aged (10–12 mo), and aged (22–24 mo) mice, we observed that aged C57Bl/6 male mice
Keywords:
make more errors on the challenging beam task and take fewer hind limb steps as compared to young and
Aging
Mice middle-aged mice. Body weight and food intake were also measured to determine if these parameters
Motor were confounding factors in the interpretation of the behavioral data. Increases in body weight and food
Behavior consumption were not observed in the oldest group that made the most errors. Together these data
indicate that aged C57BL/6 mice display age-related motor deficits similar to those seen in humans and
are an appropriate model of motor deficits that occur with age.
© 2014 Elsevier B.V. All rights reserved.

1. Introduction may lead to long-term disability, pain, and untimely death [5]. For
example, falls are the leading cause of unintentional injury and
The percentage of people aged 65 years and older has increased injury-related death in the elderly. Subsequently, this increase in
from 12.4% of the United States (US) population in 2000 to 13.0% falls leads to an increase in the risk of fractures, with 20% of falls
of the US population in 2010, with a projected increase to 20.2% requiring admission to the hospital [6].
by 2050 [1]. Moreover, the number of people aged 65 and older Currently, this age-related decline in motor skills is not treated
is projected to more than double from 41.4 million in 2011 to 92 clinically unless the loss of motor coordination is linked with dis-
million in 2060 [1]. The significance of this doubling of the aged eases, such as PD or AD. Therefore, a further investigation into the
population is that the incidence of age-related injuries and diseases cause and/or treatment of the decline in motor coordination with
will increase leading to a concomitant increase in health care costs. age is necessary. In order to study motor deficits that occur with-
Aging is associated with a decline in motor coordination and out a disease state there is a need for a model of normal aging.
increases in cognitive deficits, chronic illnesses, and diseases, such Rodent models have been widely used to investigate age-related
as Parkinson’s disease (PD) or Alzheimer’s disease (AD) [2], [3], physiological changes, including alterations in body composition
and [4]. Of these ailments, a loss in motor coordination becomes and sensorimotor behaviors [7], [8], [9], and [10]. However, many
increasingly debilitating with age as the risk of injury from falling of the studies on aging lack a middle-aged group. The purpose of
this study was to examine the appropriateness of the use of male
C57Bl/6 mice as a model for age-related motor decline. In order to
∗ Corresponding author at: Department of Pharmacology, 455 Mellon Hall, examine alterations that may occur across the lifespan of C57Bl/6
Duquesne University, United States. Tel.: +1 412 396 6358; fax: +1 412 396 4660. mice, we included three age groups to mimic young, middle, and
E-mail address: cavanaughj@duq.edu (J.E. Cavanaugh). old age. We hypothesized that the male C57Bl/6 mice will be an

http://dx.doi.org/10.1016/j.bbr.2014.03.032
0166-4328/© 2014 Elsevier B.V. All rights reserved.
120 E.N. Allen, J.E. Cavanaugh / Behavioural Brain Research 267 (2014) 119–125

appropriate model of age-related motor decline with the oldest 2.4. Statistical analysis
animals exhibited the greatest loss of motor coordination. We also
measured food intake and body weight to exclude these parameters All statistical tests were performed using Graph Pad Prism 5.
as confounding factors for our data. The data were analyzed using a one or two-way ANOVA (p < 0.05)
followed by post hoc comparison using an uncorrected Fisher’s LSD
2. Methods test. The data were analyzed using an unpaired t-test with a 95%
confidence interval.
2.1. Animals
3. Results
Young (2 mo), middle-aged (10 mo), and old (22 mo), male,
C57BL/6 mice were obtained from the National Institute on Aging 3.1. Assessment of age-related motor deficits by the spontaneous
(NIA) aging colony (Charles River) and were housed singly. All ani- activity task
mals were maintained on a 12-h light:12-h dark cycle. Water and
food were supplied ad libitum until the study began. Food supplied In the spontaneous activity test mice are placed in a Plexiglas
was Purina LabDiet® 5015 (St. Louis, MO). All procedures were con- cylinder for 3 min. Each mouse was analyzed for the number of
ducted in accordance with the guidelines for the NIH Care and Use times a rearing movement is made, the number of forelimb and
of Laboratory Animals and approved by the Duquesne University hindlimb steps, and amount of time spent grooming. Aged mice
Institutional Animal Care and Use Committee. took significantly less hind limb steps than young or middle aged
mice (F2,22 = 2.6194, p ≤ 0.05; Fig. 1C). However, there was not a
significant age-related difference in rears, forelimb steps, or time
2.2. Behavioral analysis
spent grooming (Fig. 1A, B and D).
2.2.1. Cylinder test for spontaneous activity
3.2. Assessment of age-related motor coordination by the
Animals were tested for spontaneous movement as previously
challenge beam task
described [11] and [12]. For these experiments, a small, transparent,
Plexiglas cylinder (height 15.5 cm, diameter 12.7 cm) was placed
The challenge beam task uses a beam that gets progressively
on a piece of glass that was elevated by four pedestals (6 in.). A
narrower to test for skilled walking and overall motor coordi-
mirror was placed underneath the glass and angled towards the
nation as a mouse traverses towards its home cage. An ANOVA
video camera so that the mice could be viewed from below. Ani-
indicated a significant effect of age on the challenge beam task
mals were videotaped in the cylinder for 3 min. Videos were rated
(F2,112 = 5.525, p ≤ 0.001) such that aged (22 mo) C57Bl/6 male
in slow motion for forelimb steps, hind-limb steps, rears, and time
mice made significantly more errors than young animals (2 mo;
spent grooming by an investigator blinded to the treatments. A step
Fig. 2A). Interestingly, post hoc analysis revealed that aged mice also
was counted when the animal moved both fore paws or hind paws
made more errors than middle-aged mice (10 mo; p-value = 0.053,
across the glass and a rearing movement was considered as a verti-
Fig. 2A). In order to determine if the errors made were altered by
cal movement with both front paws leaving the glass floor. Values
other parameters of the challenging beam task, we determined the
were calculated as the mean ± SEM.
number of steps, the time to transverse the beam, and the errors
per segment of the beam. Young and old animals took a similar
2.2.2. Challenging beam test for motor coordination number of steps when crossing the beam. In contrast, middle-
Animals were tested on the challenging beam to assess motor aged mice took fewer steps traversing the beam than young mice
performance as previously described [13], [14], and [15]. Briefly, (F2,112 = 5.720, p ≤ 0.001 (Fig. 2B). Even with this decline in the num-
the mice were trained to traverse an elevated beam (4 segments, ber of steps in the middle-aged group, analysis of the numbers of
25 cm long × 3.5 cm at the widest segment, narrowing to 0.5 cm errors per step (errors/step) revealed a significant increase in this
in 1 cm increments) and enter their home cage on two separate measurement in the old age group (data not shown). Although we
days. One week after training, wire mesh with 1 cm2 openings was hypothesized the old mice would take longer to cross the beam to
placed over the beam segments and the animals were videotaped their home cage, our data suggest that the time to cross the beam
and timed for 3 traversals. This initial testing was considered week is not significantly altered with age (Fig. 2C).
1. Animals were tested again mid-way through the study (week Since the challenge beam becomes more difficult as the animal
4) and at the end (week 8). Therefore, each animal was tested on progress (3.5 cm to 0.5 cm × 1 cm increments; 1 m total length), we
3 separate days (weeks 1, 4 and 8). Videotapes were viewed and examined the number of errors the mice made on each segment of
rated in slow motion for the time to transverse and number of steps the beam to determine if the age-related changes in motor coordi-
and errors by an investigator blinded to the treatment group. An nation are dependent on the width of the beam. In each age group,
error was counted when any paw passed through the mesh grid the mice make the most errors on the narrowest portion of the
and broke the plane of the grid or the paw slipped off the side of beam (F3,444 = 78.79, p ≤ 0.0001; Fig. 2D). Importantly, middle-aged
the mesh grid. Each paw was counted as a separate error, i.e., if two and old mice made significantly more errors than young mice on
paws slipped through the grid at one time they were considered the narrowest part of the beam (F2,444 = 12.23, p ≤ 0.0001; Fig. 2D).
two errors. Time to cross the beam was considered from the time
the mouse started forward on the beam to the time it took the 3.3. Age-related changes in body weight and food intake
mouse to reach the end of the beam. The means of three trials were
used for analyses. As weight of the animals may affect their ability to cross the
beam such that a larger body size may lead to decreased mobility
2.3. Food intake and weight measurement or more difficulty balancing on the beam, body weights and food
intake were recorded over the eight week testing period. Young
Animals were given a fresh supply of 6 g of food daily. Food was and old mice did not gain a significant amount of weight at any
placed on the floor of the cage to provide adequate and equal access time during the eight weeks as compared to the first day of the
for all age groups. Body weight (in grams) and food intake (in grams) study (Fig. 3A). However, middle-aged mice gained a significant
were measured each morning when a new supply of food was given. amount of weight throughout the duration of the study as revealed
E.N. Allen, J.E. Cavanaugh / Behavioural Brain Research 267 (2014) 119–125 121

A B
20 150

F o r e lim p s t e p s
15
100
R e a r in g

10

50
5

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T im e s p e n t g r o o m in g (s e c o n d s )
100 8

80
H in d lim b s te p s

6
**
60
4
40

2
20

0 0
ld
g

le

ld
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O
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Fig. 1. Aged mice take fewer hind limb steps than young mice. Results are presented as mean ± SEM. n = 37 young, 40 middle-aged, and 38 old animals. Spontaneous activity
was measured by counting rears, forelimb and hind limb steps, and grooming. Age did not alter the average amount of rears (A) or forelimb steps (B) or time spent grooming
(D) made by the mice. However, old mice made significantly less hind limb steps than young animals (C, *p ≤ 0.05 vs young). Data was analyzed using a one-way ANOVA
followed by post hoc analysis using an uncorrected Fisher’s LSD test.

by post hoc analysis (Fig. 3A). Moreover, middle-aged mice con- 4. Discussion
sumed more of the daily diet (F2.2,8.9 = 3.689, p = 0.06; Fig. 3B) and
young mice weighed significantly less than middle-aged or old ani- A number of deficits, both behavioral and neurological, have
mals throughout the 8 weeks of the study (F2,88 = 254.9, p ≤ 0.001; been shown to increase with age. Due to the rise in the aging
Fig. 3A). Moreover, post hoc analysis revealed that middle-aged population and a concurrent increase in age-related health prob-
mice weighed significantly more than old mice over the eight-week lems, such as a decrease in motor coordination, there is a need for
testing period. an appropriate model of motor aging. In the present study, age-
related differences in motor performance were assessed in young,
middle-aged, and old, male, C57BL/6 mice using the cylinder and
3.4. Assessment of age-related motor coordination by the challenging beam behavioral tests to measure spontaneous activity
challenge beam task at 4 and 8 weeks after the initial test and balance and coordination, respectively. Similar to other studies
[16], [17], [18] and [19], age-related decreases in motor function
In order to determine whether the motor deficits observed ini- were observed. However, a unique aspect of the current study is
tially in the aged group would progress further, mice were retested the inclusion of three age groups, particularly a middle-aged group.
on the challenge beam four and eight weeks after the initial test Many previous behavioral studies include only an old aged group
day. At four weeks, old mice made significantly more errors than and do not make comparisons across different ages. Interestingly, in
young (p ≤ 0.001) and middle-aged (p ≤ 0.01) mice on the challenge the current study, a progressive loss of motor coordination with age
beam (F2,25 = 8.811, p ≤ 0.01; Fig. 4A). At eight weeks, only aged mice was observed, with deficits in motor function as measured by both
made more errors than young mice (p = 0.06; Fig. 4B) and the differ- the cylinder and challenging beam tasks starting at middle-aged.
ence between middle-aged and old mice was no longer significant. This loss of motor coordination in the middle-aged animals under-
This is due to the fact that middle-aged mice made more errors scores the importance of including representative groups across the
at week eight than week four (5.6 and 3.8, respectively; Fig. 5), lifespan. To further validate our animal and behavioral paradigms,
while aged mice made approximately the same number of errors confounding factors such as number of steps and time taken to cross
at weeks four and eight (6.9 and 7.0, respectively). Surprisingly, the beam and weight and food intake were measured. Although dif-
post hoc analysis revealed that middle-aged mice make more errors ferences in the number of steps taken to cross the beam, weight and
at week eight than either week one or four; p ≤ 0.05 and p = 0.06, food intake were observed, these differences did not correlate with
respectively (Fig. 5). alterations in motor performance.
122 E.N. Allen, J.E. Cavanaugh / Behavioural Brain Research 267 (2014) 119–125

.0 5 3 25
A 6 B **
**
20

4
15

S te p s
E rro rs

10
2

0 0
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M
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C 25
10
D **** S egm ent 1
20
T im e ( s e c o n d s )

8 S egm ent 2
# o f e rro rs
** S egm ent 3
15
6 S egm ent 4

10 4

5 2

0 0

ld
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Fig. 2. Aged mice make more errors than young- and middle-aged mice. Results are presented as mean ± SEM. n = 37 young, 40 middle-aged, and 38 old animals. C57Bl/6
male mice display age-related motor deficits as measured by the challenge beam task. (A) Old mice made significantly more errors than young mice (**p ≤ 0.01) and more
errors than middle-aged mice (p = 0.053). (B) Middle-aged mice took significantly less steps while traversing the challenge beam than young mice (**p ≤ 0.001). The challenge
beam narrows as the mouse traverses towards its home cage. (C) Middle-aged and old mice make significantly more errors on the narrowest segment of the beam than
young mice (**p < 0.01 and ****p ≤ 0.0001). Data was analyzed using a one or two-way ANOVA followed by post hoc analysis with an uncorrected Fisher’s LSD test.

Using the cylinder test as a measure of spontaneous activity, no While the cylinder test provides a measure of spontaneous activ-
significant differences in the number of rearing movements, fore- ity and motor function in general, the challenging beam task has
limb steps or time spent grooming were noted in the three age been used as a highly sensitive measure of motor coordination [11],
groups. In contrast, the number of hind limb steps was significantly [13], [23] and [25]. In the current study, an age-related decline in
decreased in the old animals as compared to the young animals. motor coordination, as measured by the number of errors made
Sustained deficits in forelimb and hind limb functions have been on the challenging beam, was noted. Using the challenging beam,
noted in aged Wistar rats following focal cerebral ischemia induced Fleming et al. [23], [25] noted similar declines in motor coor-
by unilateral endothelin-1 infusions into the sensorimotor cortex dination in transgenic mice expressing human alpha-synuclein.
[20]. Others have reported improved hind limb function follow- Moreover, an increase in the number of errors on the beam was
ing electroconvulsive shock-enhanced striatal dopamine receptor noted in mice unilaterally lesioned with the dopaminergic toxin,
1 (D1) and D3 binding [21] or L-DOPA treatment following mid- 6-hydroxydopmaine (6-OHDA; [11]). Together, these data again
dle cerebral artery occlusion in rats [22]. These data suggest that suggest that alterations in the nigrostriatal system may underlie
an age-related decrease in the number of hind limb steps may the behavioral deficits in motor function noted with age.
be due to functional declines in the sensorimotor cortex and/or In our study, young and old animals took the same number
striatal dopamine receptors with age. The involvement of the of steps when crossing the challenging beam, while middle-aged
nigrostriatal dopaminergic system in the loss of hind limb motor mice took fewer steps than aged mice. However, the time that it
activity is further supported in a study by Fleming et al. [23], in took each group to cross the beam was not significantly different
which a decrease in the number of hind limb steps was noted in between the three age groups. Despite the fact that middle-aged
transgenic mice expressing human alpha-synuclein leading to dys- animals took fewer steps to cross the beam, the number of errors
function of the dopaminergic system. Interestingly, in a previous progressively increases with age. Also, with further analysis, old
study, we did not observe a decline in striatal dopamine, dopamine mice made significantly more errors per step (errors/step) than
metabolites, tyrosine hydroxylase, or the dopamine transporter young and middle-aged mice (data not shown). Therefore, our data
with age [24]. Therefore, age-related declines in striatal dopami- suggest that the number of errors measured on the challenging
nergic function that result in a loss of motor function may be beam reflects a decline in motor performance with age that was
limited to alterations in the D1 and D3 receptors in C57Bl/6 male not confounded by alterations in the number of steps or time taken
mice. to cross the beam.
E.N. Allen, J.E. Cavanaugh / Behavioural Brain Research 267 (2014) 119–125 123

Young
A B
6
50 Young
****
** *** M id d le

F o o d in t a k e ( g )
* * O ld
40 4
W e ig h t ( g )

30

2
20

10 0
0

7
1

5
y

y
a

y
a

a
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a
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D
D

D
C D
M id d l e - a g e d O ld
6 6
****
** * **
F o o d in t a k e ( g )

F o o d in t a k e ( g )
*
4 4

2 2

0 0
0

8
5

7
1

5
y

y
a

a
y

y
a

a
D

D
D

D
Fig. 3. Middle-aged mice weighed and ate more than young and aged mice over the course of the 8 week study. Results are presented as a mean ± SEM. n = 5. (A) Body
weight (in grams) was measured daily. Middle aged mice gained a significant amount of weight over an eight week period (*p < 0.05, **p < 0.01, ***p < 0.001, ****p < 0.0001 vs
middle-aged day 0). (B–D) Food intake was monitored daily by administration of 6 g/day and weight of food (in grams). Middle-aged mice ate significantly more throughout
the eight-week study as compared to the beginning (*p < 0.05, **p < 0.01, ***p ≤ 0.001 vs middle–aged day 0). Data was analyzed using a repeated measures two-way ANOVA
followed by post hoc analysis with an uncorrected Fisher’s LSD test.

A W eek 4 B W eek 8
10 10
***

.0 6
**

8 8
E rro rs

6 6
E rro rs

4 4

2 2

0 0
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ld
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Fig. 4. Aged mice make more errors than both young and middle-aged mice at 4 weeks following the first behavioral test but only make more errors than young mice at 8
weeks. Results are presented as mean ± SEM. n = 8–10. Mice were retested on the challenge beam task after 4 and 8 weeks. (A) Aged mice made significantly more errors
than young (***p ≤ 0.001) and middle-aged (**p ≤ 0.01) mice. (B) After 8 weeks, there is no difference in errors between middle-aged and old mice. However, old mice made
more errors than young mice (p = 0.06). Data was analyzed using a two-way ANOVA followed by post hoc analysis with an uncorrected Fisher’s LSD test.
124 E.N. Allen, J.E. Cavanaugh / Behavioural Brain Research 267 (2014) 119–125

8 [35]. However, other studies which included young, middle-aged,


.0 5 and old, male, C57Bl/6 mice reported increases in body weight up
*
to 20–24 months of age [36] and [37]. This may be due to a dif-
6 ference in diet fed to the mice as Blackwell and colleagues used an
NIH-31 diet while the current study used Purina Standard LabDiet®
E rro rs

5015. Along with the increase in fat mass with age, a decline in
4 lean muscle mass has been noted leading to muscle weakness [32],
[33] and [34]. Therefore, it is plausible that muscle weakness con-
tributed to the increase in errors. However, we did not see an overall
2 decline in spontaneous activity nor did we observe a change in body
composition throughout the study.
The increase in body weight in the middle-aged mice is accom-
0 panied by a significant increase in food consumption by this
group throughout the 8 week study. One caveat when interpre-
1

8
ting the food intake data, however, is that it has been reported
k

k
e

e
e

e that the amount of food spilled during consumption is significantly


W

increased by 20–21 months of age in C57BL/6 mice [37]. There-


Fig. 5. The number of errors made by middle-aged mice increases over eight weeks. fore, the food intake measured for the old animals in the current
Results are presented as the mean ± SEM. n = 10. The number of errors was averaged
study may be higher than they actually consumed. Nonetheless,
from three trials for each group. At weeks 1 and 4, middle-aged mice made an aver-
age of 3.8 errors, which increases to an average of 5.6 errors by week 8. Middle-aged body weights are accurate measurements, and as aged mice made
mice made significantly more errors at week 8 than week 1(*p < 0.05) and week 4 significantly more errors on the challenge beam task as compared
(p = 0.06). Data was analyzed using a repeated measures one-way ANOVA followed to young and middle-aged mice, weight does not appear to be a
by post hoc analysis with an uncorrected Fisher’s test. confounding factor in the current study.
An important and novel aspect of this study was to the repeated
Another factor that must be taken into consideration when using examination of the effect of age on motor coordination over an 8
the challenging beam test is that there is a component of motor week period. The challenge beam task is normally used only once
learning and exercise associated with this task. Motor learning and before animals are removed from a study [13], [14] and [15]. How-
exercise have been shown to delay age-related cognitive deficits ever, in the current study, the mice were tested at baseline and
[26] and increase levels of brain derived neurotrophic factor (BDNF) four and eight weeks following the start of the study. The purpose of
[27] and [28], an established neurotrophic factor for dopaminergic this design was to determine if age-related motor deficits increased
neurons [29] and [30]. However, our aged animals did not show over time and to validate the repeated use of the challenging beam
improvement in the challenge beam task even at week 8. Therefore, behavioral test. Old animals made significantly more errors than
the time intervals at which they were tested may be sufficient to young and middle-age animals at week 4, but by week 8 old ani-
decrease or eliminate the impact of motor learning and exercise on mals only made significantly more errors than the young age group.
the behavioral task. Most interesting is a comparison of the number of errors made by
An additional caveat of the current study design is that the mice the middle-aged group that entered the study at 10 months of age
were weighed and, therefore, handled daily. The handling of the and were last tested at 12 months of age. By week 8 of the study,
mice daily could be considered a form of environmental enrich- when the middle-aged mice were 12 months old, there was a sig-
ment. In fact, O’Callaghan et al. noted that animal handling had a nificant increase in the number of errors made by the middle-aged
significant effect on learning as measured by the water maze task mice as compared to the same age group at the start of the study
[31]. Therefore, it is possible that our daily handling of the mice and week 4. Moreover, by week 8, old mice no longer made signifi-
may have improved their motor learning and thereby influenced cantly more errors than the middle-aged group. These data suggests
the behavioral results. However, all animals in each age group were that a decline in motor coordination may begin to occur between
handled at the same time of day and to an equal extent. Further- 10 and 12 months in this animal model. In contrast to the present
more, old mice made significantly more errors than young mice study, most behavioral studies using aged animal models do not
even at 8 weeks following the initiation of the studies suggesting include a middle-aged group or comparisons between age groups.
that the handling of the animals for this time period did not increase As behavioral changes were noted at week 8 in the middle-aged
their motor learning. animals, the current data underscore the importance of including
In addition to the potentially confounding behavioral and learn- this middle-aged group. Moreover, these data suggest that treat-
ing parameters for the challenging beam task, significant weight ments aimed at alleviating age-related motor decline may be more
gain could cause a decrease in mobility and contribute to the age- effective if started at middle-age rather than later in life.
related increases in the number of errors and loss of hind limb In conclusion, there is a progressive age-related decline in motor
function noted in the C57Bl/6 mice. Therefore, body weights and performance in male C57Bl/6 mice with the start of the decline
food intake were recorded over the eight-week testing period. beginning at middle-age. The loss of motor function with age
These measurements were also used to insure that animals were occurs without a significant contribution of possibly confound-
receiving comparable amounts of food and stave off weight loss. ing factors such as the number of steps, time to cross the beam,
Metabolism is known to slow with age, in part, due to an age-related weight, or food intake. Finally, the loss of motor coordination
increase in fat mass [32], [33], and [34]. Therefore, we expected that in the old mice is evident up to eight weeks following the ini-
the animals in the old age group would weigh more than the young tial behavioral test. Together these data suggest that there is a
and middle-aged mice. Surprisingly, middle-aged, but not old, ani- chronic loss of motor function with age that can be measured
mals gained a significant amount of weight and weighed more than using the C57Bl/6 mice as an animal model and the cylinder and
young and old mice throughout the course of the study. The data challenging beam tasks for behavioral measurements. Moreover,
from the current study on body weight in the young and old groups the importance of including a middle-aged group is highlighted
are in agreement with a previous study in which the weights of by the observation that motor deficits begin to appear in this age
5-month- and 17-month-old mice were not significantly different group.
E.N. Allen, J.E. Cavanaugh / Behavioural Brain Research 267 (2014) 119–125 125

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