Scand J Med Sci Sports 2016: 26: 29–40
doi: 10.1111/sms.12400
Stress responses to short-term intensified and reduced training in
competitive weightlifters
A. G. Storey1, N. P. Birch2, V. Fan3, H. K. Smith1
1
Department of Sport and Exercise Science, The University of Auckland, Auckland, New Zealand, 2School of Biological Sciences,
Centre for Brain Research and Brain Research New Zealand, The University of Auckland, Auckland, New Zealand, 3Bioinformatics
Institute, The University of Auckland, Auckland, New Zealand
Corresponding author: Heather K. Smith, PhD, Exercise and Muscle Physiology Laboratory, Department of Sport and Exercise
Science, The University of Auckland, Private Bag 92019, Auckland Mail Centre, Auckland 1142, New Zealand. Tel: +64 9 373 7599,
Fax: +64 9 373 7043, E-mail: h.smith@auckland.ac.nz
Accepted for publication 4 December 2014
We sought to identify and evaluate the tolerance to, and
consequences of, short-term variations in training load in
competitive weightlifters. Seven international-level lifters
performed 1 week of initial training followed by 2 weeks
of intensified (INT: +100%, 36.5 ± 11.3 × 103 kg/week)
and 1 week of subsequently reduced (RED: −25%) train-
ing within their annual program. After INT, but not
RED, 90 min of weightlifting increased mRNA levels of
chemokine (C-C motif) ligand 4 (CCL4), chemokine
(C-X-C motif) receptor 4 (CXCR4) and cellular stress-
associated DNA-damage-inducible transcript 4 (DDIT4)
in peripheral blood mononuclear cells by 40–240%.
Resting- and weightlifting-induced changes in plasma
Systematic variations in exercise training intensity and
volume, or training load, are hallmarks of training pro-
grams prescribed to improve competitive athletic perfor-
mance. In competitive weightlifting, characteristic
variations in training load include periodic, short-term
(1–3 week) increases or “overload” and subsequent
decreases or “recovery” within phases of a longer-term
training program (Gonzalez-Badillo et al., 2006; Haff
et al., 2008; Pistilli et al., 2008). The intention of these
periods of intensified (INT) and reduced (RED) training
is improved performance during a subsequent competi-
tive phase. However, despite being common practice in
weightlifting, evidence of the physiologic, psychologic,
and performance consequences of such variations in
training load is limited (Warren et al., 1992; Fry et al.,
1993, 1994, 2000; Stone & Fry, 1998). These studies
have described variations in serum hormones (e.g., tes-
tosterone, cortisol) and changes in neuromuscular
function that only partially explain the corresponding
expected or actual changes in performance.
Our review of the performance, training, and physi-
ologic aspects of weightlifting suggested that numerous
diverse and complex, system-wide responses and
adaptations contribute, collectively, to competitive
© 2015 John Wiley & Sons A/S.
Published by John Wiley & Sons Ltd
protein carbonyls, indicative of oxidative stress, but not
pro-inflammatory CCL4 concentrations differed between
INT and RED. Symptoms of stress (Daily Analysis of Life
Demands of Athletes questionnaire) were reported as
worse than normal more frequently during INT and RED
than initial training. Global (negative) mood state
increased during INT and declined during RED.
Maximal snatch (−4.3 ± 3.7%) and vertical jump
(−7.2 ± 6.5%), but not clean and jerk, were reduced after
INT and restored after RED. Chemokine signaling may
thus be part of the stress response to intense weightlifting
and short-term reductions in training load support recov-
ery from periodic INT training in weightlifters.
performance (Storey & Smith, 2012). Further to the avail-
able evidence, we conjectured that non-endocrine, sys-
temic (blood-borne) cells and molecules might act as
inter-organ communicators in the adaptation of tissues
and mediate the anticipated, and often rapid, functional
adaptations (changes in performance) to short-term varia-
tions in training load. To this end, peripheral blood mono-
nuclear cells (PBMC) presented an accessible and
potentially important source of insight (Fehrenbach,
2007).
Furthermore, we expected that any differences in sys-
temic responses between INT or RED training loads
would be most evident during the early recovery after
training sessions. This concept is supported by evidence
of weightlifting-induced changes in testosterone that
differed after a period of INT training (Fry et al., 1994),
whereas long-term resting testosterone levels in
weightlifters remain relatively stable (Häkkinen et al.,
1989).
We therefore took a discovery approach, using full
genome microarray analyses of PBMC, to first identify
weightlifting-responsive genes in a subset of partici-
pants. Differential expression, pathway analyses, and
evidence from the literature suggesting plausible roles in
29
Storey et al.
relation to athletic training, led to our selection and
extended analysis of the genes CCL4, CXCR4, and
DDIT4.
These three genes respond to, and have functions in,
cellular stress, damage, and inflammation (Menten et al.,
2002). The gene CCL4 encodes the chemokine CCL4,
also known as macrophage inflammatory protein-1β.
The chemokine is secreted from mononuclear cells and
has pro-inflammatory and regenerative effects in tissues,
including skeletal muscle (Warren et al., 2004; Yahiaoui
et al., 2008). An increased expression of CCL4 in skel-
etal muscle 2 h after resistance exercise in older adults
has also been reported (Mathers et al., 2012). The second
gene, CXCR4, has numerous diverse functions, being
expressed by several cell types including lymphocytes
and monocytes. Circulating CXCR4-positive cells are
recruited by the chemokine (C-C motif) ligand
12/stromal cell-derived factor-1 (CXCL12/SDF-1) to
injured tissues, including skeletal muscle (Kucia et al.,
2004; Ratajczak et al., 2006). The third gene, DDIT4, is
upregulated during cellular stress, including DNA
damage and hypoxia (Drummond et al., 2008). DDIT4
inhibits mTORC1, a major regulator of protein synthesis
(+) and autophagy (−), via the TSC1–TSC2 complex
(Drummond et al., 2009; Ma & Blenis, 2009). Its expres-
sion has been reported to decline in untrained human
skeletal muscle after resistance exercise and essential
amino acid ingestion (Drummond et al., 2009).
Carbonylated proteins, products of protein oxidation,
are indicative of oxidative stress in skeletal muscle
(Margonis et al., 2007; Veskoukis et al., 2009). The
plasma concentration of protein carbonyls has been
shown to increase in response to high-volume, high-
intensity eccentric exercise protocols known to induce
muscle damage (Lee et al., 2002) and with high-intensity
resistance exercise in trained adults (Hudson et al., 2008).
We therefore examined the mRNA levels of CCL4,
CXCR4 and DDIT4 in PBMC and plasma CCL4 and
protein carbonyl concentration as systemic indices of
cellular stress, damage, and inflammation in response to
weightlifting after short-term INT and subsequently RED
training. In conjunction with the physiologic responses,
we evaluated the psychologic and performance conse-
quences of the variations in training load, in competitive
weightlifters. We anticipated that a weightlifting session
would increase the systemic indices more after INT train-
ing than after subsequently RED training. In parallel, we
expected the psychologic profile and weightlifting perfor-
mance of the athletes to decline with the INT training and
subsequently improve with RED training.
Methods
Experimental design
This study was designed to compare resting- and training session-
induced systemic responses and changes in performance with
short-term INT and subsequently RED training in competitive
weightlifters (Fig. 1). The experimental approach and methods
30
were guided by an accurate representation of, and compatibility
with, athlete training practices. Participants completed a 4-week-
long training cycle within a pre-competitive period of their annual
program. Structured preparatory training was performed leading
up to the study. The training load prescribed during the second and
third weeks (INT) was approximately double that during the initial
week. It was then reduced by approximately 25% for the fourth
week (RED). Central to this study was the identification and com-
parison of systemic indices of cellular stress, damage and inflam-
mation in response to the same high-intensity, 90-min-long
weightlifting training session, performed under the same condi-
tions, at the end of INT and again after RED. During INT and
RED, performance was evaluated by snatch and clean and jerk
achieved during simulated weightlifting competition and maximal
vertical jump height. Two questionnaires were administered to
evaluate symptoms of stress and mood state of the athletes
throughout the study (Fig. 1).
Subjects
Seven competitive weightlifters [4 men, 3 women; mean ± stan-
dard deviation (SD), range: body mass 80.4 ± 18.6, 54.5–
104.2 kg; height 165.4 ± 9.1, 150–177 cm; age 22.9 ± 4.3, 19–30
years] volunteered to participate in this study. All had International
competitive lifting experience, ranging from Oceania Champion-
ships to the Olympic Games. As a requirement of participation, the
lifters were free of injury and were not using anti-inflammatory
medication or any performance-enhancing and/or banned sub-
stances [World Anti-Doping Agency (WADA), The 2010 Prohib-
ited List]. The participants gave written informed consent before
commencing with the study. The study was approved by The
University of Auckland Human Participants Ethics Committee.
Training program
The frequency, intensity, and volume of weightlifting performed
during the initial, INT, and RED training weeks are shown in
Table 1. The training comprised the competitive lifts (snatch, clean
and jerk), related lifts (power snatch and power clean), and supple-
mentary exercises (front squats, back squats, clean pulls, snatch
pulls). The intensities (kg) for each set of each exercise were
prescribed relative to each athlete’s recent best performances
(1-repetition maximum) in competition or in training. The rest
intervals between sets and exercises ranged from approximately 1
to 5 min, as determined by the athletes’ preparedness and the
coach’s discretion, such that the duration of each training session
was 90–100 min. All sessions were prescribed and administered by
one of the investigators in the athletes’habitual training facility. The
training loads shown in Table 1 were calculated as those completed,
using the weights (bar and plates) lifted (kg) during training, for all
exercises.
Responses to weightlifting training after INT and
RED training
To allow for the acute responses to weightlifting after INT and
RED to be compared, the final training session of INT and RED
followed the same prescription. A standard warm+up was followed
by six to eight sets of one to three repetitions, of progressively
increasing intensity, of power snatch, power clean, and back squat
exercises. The final two sets for each exercise were performed at
90% of maximum for one (power snatch and power clean) or three
(back squat) repetitions. For these sessions, the rest intervals
between sets and exercises were timed such that the duration of
each session was consistently 90 min. In addition, before the ses-
sions at the end of the initial, INT, and RED trainings, participants
 Stress responses in competitive weightlifters
17:30 →

W W W W W SC W W W SC W W W SC
↑ 16:00 →
D D D D D D D D D D D D D D D D D D D D D D D D D D D D
TIME OF DAY

13:30 → B: M, PCR B: PCR

CCL4, PC CCL4, PC

SM SM
↑ 10:30 → B: CCL4, PC B: CCL4, PC

W W W W
TIME OF DAY

W W W W W W W W W W W W W W W W
VJ VJ VJ VJ
09:00 →

08:00 → SM SM SM
B: PCR B: M, PCR B: PCR
↑ CCL4, PC CCL4, PC
P P P P P P P P P P P P P P P P P P P P P
DAY → M T W T F S S M T W T F S S M T W T F S S M T W T F S S
WEEK → 1 2 3 4
TRAINING → INITIAL INTENSIFIED REDUCED

W Weightlifting training session B Venous blood sample

SC Maximal snatch and clean and jerk Assays:
VJ Vertical jump M Microarray
SM Standardized meal PCR qRT-PCR: CCL4, CXCR4, DDIT4 mRNA
D DALDA questionnaire CCL4 CCL4 protein
P POMS questionnaire PC Protein carbonyls

Fig. 1. Experimental design. Weightlifting training sessions, maximal snatch, clean and jerk, and vertical jump performance tests and
psychological questionnaires completed over 4 weeks of training within a pre-competitive phase of the annual program. The same
performance testing protocols, training sessions, and blood sampling intervals were used during the intensified training and reduced
training (bold boxes). DALDA, Daily Analysis of Life Demands of Athletes; POMS, Profile of Mood States questionnaire; qRT-PCR,
reverse transcription and real-time polymerase chain reaction.

Table 1. Training loads during initial, intensified, and reduced training weeks in weightlifters

Training Initial Intensified Reduced

Week 1 2 3 4

Frequency
Sessions per week 6 10 10 9
Sessions per week relative to initial (%) 100 167 167 150
Intensity
Peak (%1RM) 85 95 97 88
Peak relative to initial (%) 100 112 114 104
Mean (kg) 77 ± 22 86 ± 24 89 ± 25 80 ± 24
Mean relative to initial (%) 100 112 115 104
Load / Volume
Total repetitions 239 ± 14 414 ± 20 414 ± 20 336 ± 26
Total repetitions relative to initial (%) 100 173 173 141
Total (×103 kg) 18.5 ± 6.0 35.9 ± 11.5 37.1 ± 11.9 27.0 ± 8.4
Total relative to initial (%) 100 194 200 146

Values are means ± standard deviation.

1RM, one repetition maximum.

arrived after an overnight fast, provided a venous blood sample,

and were given a standard meal (48 kJ/kg body mass, with
56% from carbohydrate, 33% from protein, and 11% from
fat) to consume. They then rested for 1 h before training. After
training, participants provided another blood sample and were
given a second meal (as previous). Three hours after completion
of the training session, a third blood sample was collected
(Fig. 1).
RNA isolation from PBMC
Venous blood samples were drawn into BD Vacutainer™ CPT™
Sodium Citrate Tubes (Becton Dickinson, Heidelberg, Germany).
The blood samples were processed within 15 min to isolate mono-
nuclear cells suitable for gene expression profiling by microarray
analyses (Affymetrix, 2003; Debey et al., 2004). Plasma obtained
during the process was stored at −80 °C until further analyses. The

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Storey et al.
mononuclear cell pellet was stabilized in RNAlater™ RNA Sta-
bilization Reagent (Ambion, Inc., Austin, Texas, USA).
Total RNA was isolated and purified from PBMC using a com-
bined TRIzol reagent (Invitrogen, Karlsruhe, Germany) and mini
spin column (RNeasy® Mini Kit, Qiagen, Hilden, Germany) pro-
tocol. Total RNA concentration and purity was determined using a
Nanodrop spectrophotometer (ND-1000, Thermo Fisher Scien-
tific, Wilmington Delaware, USA). Samples with a 260/280 absor-
bance ratio ≥ 1.8 were used. The quality of RNA samples was
determined with an Agilent 2100 Bioanalyzer® and RNA6000
Nano LabChips (Agilent Technologies, Palo Alto, California,
USA). The generated RNA integrity number (RIN) of samples
used was > 7.0, with > 90% of samples having a RIN >8.0.
Microarray analyses
To identify weightlifting-responsive genes in PBMC in the first
instance, cDNA microarray analyses (Human GeneChip® Gene
1.0 ST Arrays, Affymetrix Inc., Santa Clara, CA, USA) of resting
and 3 h post-weightlifting PBMC RNA samples from the four
highest-ranked (International Weightlifting Federation Sinclair
coefficient) athletes at the end of INT (Week 3, Fig. 1) were
conducted. The RNA labeling protocol was performed according
to the GeneChip® Whole Transcript Sense Target Labeling Assay
Manual (P/N 701880, Version 4, Affymetrix). The arrays were
scanned on an Affymetrix GeneChip® Scanner 3000 7G controlled
by Command Console® Software. The data were analyzed using
the statistic programming language R. To identify outliers, raw
array intensities box plots, raw log2 probe hybridization intensities
density plots and a normalized unscaled standard error plot were
produced before the data were preprocessed and normalized using
a robust multiarray average (RMA) algorithm (Irizarry et al.,
2003). Relative log expression plots were used to check the dis-
tribution of the probe intensities after the RMA. Linear modeling
was performed on the microarray data using the R package limma.
A paired t-test was performed on the resting and 3 h post-
weightlifting pairs. An empiric Bayes method was used to mod-
erate the t-statistic (Smyth, 2004). To account for multiple testing,
a Benjamini–Hochberg correction was applied to the P-values.
The annotation of microarray probe sets having a fold change
in mRNA level from resting to 3 h post-weightlifting of ≥1.4
was used to identify differentially expressed “weightlifting-
responsive” genes. Pathway analysis was then conducted to iden-
tify networks, canonical pathways and functions associated with
the differentially expressed genes (Jiménez-Marín et al., 2009)
using commercial software (Ingenuity Systems, Redwood City,
California, USA).
Reverse transcription and real-time polymerase chain
reaction (qRT-PCR)
The microarray-identified genes CCL4, CXCR4, and DDIT4 were
quantified using qRT-PCR, in samples from all participants at rest
and 3 h after weightlifting, after both INT and RED. Each RNA
sample was treated with DNA-free™ to remove any contaminat-
ing DNA, and the DNAse after treatment, according to the kit
instructions (Ambion, Inc. and Life Technologies, Carlsbad,
California, USA). Reverse transcription was performed using
the SuperScript® VILO™ cDNA Synthesis Kit (Invitrogen, Life
Technologies).
The following TaqMan® Gene Expression Assays (Applied
Biosystems and Life Technologies) with FAM-dye labeled
probes were used: chemokine (C-C motif) ligand 4 (CCL4)
(Hs99999148_m1), chemokine (C-X-C motif) receptor 4
(CXCR4) (Hs00607978_s1), DNA-damage-inducible transcript 4
(DDIT4) (Hs01111686_g1), ribosomal protein, large P0 (RPLPO)
(Hs99999902_m1) and glyceraldehyde-3-phosphate dehydroge-
32
nase (GAPDH) (Hs99999905_m1). RPLPO and GAPDH have
previously been shown to be suitable reference genes for PBMCs
(Dheda et al., 2004). For each 20 μL singleplex reaction, 2 μL of
diluted cDNA (20 ng), 7 μL of RNase-free water, 1 μL of 20×
TaqMan® gene expression assay mix, and 10 μL of 2× TaqMan®
Universal Mastermix II (with AmpErase UNG) was used. All
samples for each participant were analyzed simultaneously (target
genes and reference genes) in triplicate using an ABI 7900HT Fast
Real-Time PCR System (Applied Biosystems). Thermal cycling
conditions were: 2 min at 50 °C and 10 min at 95 °C, followed by
40 two-step cycles including denaturation at 95 °C for 15 s and
annealing/extension at 60 °C for 1 min. No-template controls and
common samples (inter-run calibrators) were included in all
plates.
The PCR threshold cycle (Ct) data were exported into qBase+
software (Biogazelle, Belgium) for analysis and calculation of fold
changes (Hellemans et al., 2007). With this method, the target
gene expression was normalized to the geometric mean of the
reference genes and expressed relative to the resting sample from
the initial training week.
Plasma CCL4 and protein carbonyl concentration
Plasma concentrations of CCL4 were determined, at rest and at 0
and 3 h after weightlifting, at the end of INT and RED. A com-
mercial immunoassay kit (Quantikine, R&D Systems, Minneapo-
lis, Minnesota, USA) and microplate spectrophotometer (Bio-Tek
Instruments, Swedesboro, New Jersey, USA) were used for the
assay. The manufacturer’s stated sensitivity of the assay for human
plasma was less than 11.0 pg/mL with an intra-assay coefficient of
variation of 3.6%. The coefficient of determination, generated
from standards ranging in concentration from 0 to 2000 pg/mL,
was R2 = 0.99.
The concentration of carbonylated proteins was also deter-
mined in plasma samples, at the same time points as CCL4. A
commercial immunoassay kit with a sensitivity of 0.06 nmol/mg
total protein and an intra-assay coefficient of variation of 5%
(BioCell Corporation, Ltd., Auckland, New Zealand) was used.
Questionnaires
Participants completed the Daily Analysis of Life Demands of
Athletes (DALDA) questionnaire (Rushall, 1990) during the
initial, INT and RED training and the 65-item Profile of Mood
States questionnaire (POMS, McNair et al., 1971) during INT and
RED at times as shown in Fig. 1. These questionnaires, although
developed and validated in endurance sports, have proven mean-
ingful and practical tools in the monitoring of endurance and team
sport athletes and their responses to INT training (Morgan et al.,
1988; Halson et al., 2002; Nicholls et al., 2009).
The DALDA is comprised of two parts: Part A includes nine
sources of life stress and Part B includes 25 signs and symptoms of
stress. Participants responded to each item as “Worse Than
Normal,” “Normal,” or “Better Than Normal.” The POMS pro-
vided a score for the level of six mood states: tension/anxiety,
depression, anger, vigor and fatigue and confusion, at the time of
completion. A global mood state score was calculated as the sum
of the five negative mood state scores minus the score for vigor
(Morgan et al., 1987), for each day, for each athlete. The mean
weekly score was then calculated for each athlete and the mean of
all athletes calculated for each week.
Maximal snatch and clean and jerk performances
For each participant, the heaviest successful snatch and clean and
jerk lifts in official competition within 2 months of the study were
used as the initial performance levels. Maximal snatch and clean

and jerk performance was then determined after each week of INT
and after RED following weightlifting competition format and
regulations. Each participant performed a standardized general
warm-up consisting of light jogging, calisthenics and stretching,
and then a snatch specific warm-up. Participants were prescribed
eight warm-up sets progressing up to 90% of their most recent
maximal performance. In accordance with competition format,
they then attempted three lifts each separated by 2 min rest. A lift
was judged as successful, in accordance with the 2009–2012 Inter-
national Weightlifting Federation Technical and Competition
Rules, by a certified referee. Following a successful lift, the weight
of the barbell was increased by 1–5 kg to attempt and/or achieve a
maximal lift within the three attempts. After an unsuccessful
attempt, the weight remained the same for the subsequent attempt.
After the snatch, participants rested for 10 min as done in compe-
tition. A clean and jerk warm-up and maximal attempts were then
performed as per the snatch.
Maximal vertical jump
Maximal vertical jump (counter-movement with arm swing)
height was assessed using a Vertec Vertical-Jump Tester (Sports
Imports, Columbus, Ohio, USA). All participants were familiar
with the protocol as part of regular monitoring of their training.
The best of three maximal attempts was recorded. The scores were
not revealed to the athletes.
Statistics
To test for differences in mRNA levels, two factor [training load
(INT, RED) by time (resting, 3 h post-weightlifting) ] repeated-
measures analysis of variance (ANOVA) were used. Similarly, two
factor training load (INT, RED) by time (resting, 0 and 3 h post-
weightlifting) repeated-measures ANOVA were used to identify
differences in plasma protein carbonyl and CCL4 protein concen-
trations. One-way repeated-measures ANOVA were used to test
for differences in performance variables, DALDA and mean
global mood state scores, between training weeks. Where a sig-
nificant difference was determined by ANOVA, post-hoc paired
comparisons were made using the method of Student–Newman–
Keuls. All values are expressed as mean ± SD with the critical
level of significance established at P ≤ 0.05. Analyses were per-
formed using commercial software (Sigma Plot 11.0, Systat Soft-
ware, Inc., Chicago, Illinois, USA).
Results
Training program completion
The prescribed training was completed with high and
precise compliance. All athletes completed all
weightlifting sessions in full. Deviations from that pre-
scribed included only occasional repetitions of failed
attempts and minor adjustments in the weight lifted to
ensure completion of sets.
The training completed by the participants during INT
included a 73% increase in the number of repetitions and
15% increase in the mean intensity of the lifts per-
formed, the product being a 100% increase in load, rela-
tive to the initial week. During RED, participants
completed 18% fewer lifts at 9% lower intensity, result-
ing in a 27% reduction in training load, compared with
INT (Table 1).
Stress responses in competitive weightlifters
Systemic indices of cellular stress, damage
and inflammation
Gene expression in PBMCs
From the microarray analyses, we identified 202
weightlifting-responsive genes most frequently associ-
ated with the following molecular networks: (a) cell-to-
cell signaling and interaction, hematological system
development and function, immune cell trafficking; (b)
organismal survival, inflammatory response, cell-to-cell
signaling and interaction; and (c) cell cycle, cell death,
cancer. The canonical pathways and biologic functional
categories in which differentially expressed genes were
represented are shown in Table 2.
There were main effects of the weightlifting session
(time) on the mRNA levels in PBMC, determined by
qRT-PCR, of CXCR4, CCL4, and DDIT4 (Fig. 2). There
were also differences in CXCR4 mRNA between INT
and RED (training load), and interactions between time
and training load for CCL4 and DDIT4 mRNA levels
(Fig. 2).
Post-hoc paired comparisons showed that mRNA
levels of CCL4 and CXCR4 differed only after the
weightlifting session, after INT, but not RED. Similar
results were found for DDIT4, although our statistical
approach did not allow for the additional testing of an
apparent reduction in resting levels of DDIT4 mRNA
compared with the initial week (Fig. 2).
Plasma CCL4 and protein carbonyl concentration
CCL4 concentrations in plasma were low and close to
the detection level of the assay. There was no difference
in the concentration of CCL4 in plasma at rest, or at 0 or
3 h after the weightlifting session performed after INT or
RED (Fig. 3). After INT and RED, CCL4 concentration,
albeit low, decreased from resting to 3 h after the training
session to a similar extent.
The plasma protein carbonyl concentration at rest
and in response to the weightlifting session differed
between INT and RED (Fig. 4). The concentrations at
rest and directly after weightlifting were higher after
INT than RED. Moreover, the concentration declined
from rest to 3 h after the session with INT, but
increased from the lower resting levels to 3 h after the
session with RED.
Questionnaires
During INT, participants reported the signs and symp-
toms of stress (DALDA, Part B) as worse than normal
more frequently than during the initial training, an effect
that did not resolve during RED (Fig. 5). The sources of
stress (Part A) were reported as worse than normal more
often during the first week of INT than all other weeks.
Mean global mood state scores increased between the
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Storey et al.
Table 2. Top canonical pathways and biologic functions identified by pathway analysis (Ingenuity Systems, http://www.ingenuity.com) of microarray-
identified weightlifting-responsive genes

P value Number of genes relative to total in pathway (%)

Canonical pathway
Antigen presentation pathway 5.0 × 10−7 5/43 (11.6)
Allograft rejection signaling 7.7 × 10−7 5/59 (8.5)
Cytotoxic T lymphocyte-mediated apoptosis of target cells 1.3 × 10−6 5/52 (9.6)
OX40 signaling pathway 2.3 × 10−6 5/61 (8.2)
Communication between innate and adaptive immune cells 1.8 × 10−5 5/93 (5.4)
Number of genes

Functional category/biologic processes

Diseases and disorders
Dermatologic diseases and conditions ≤4.2 × 10−2 11
Immunologic disease ≤4.1 × 10−2 19
Inflammatory disease ≤4.1 × 10−2 23
Inflammatory response ≤4.9 × 10−2 12
Genetic disorder ≤3.4 × 10−2 19
Molecular and cellular functions
Protein synthesis ≤1.1 × 10−2 10
Cell signaling ≤4.6 × 10−2 8
DNA replication,recombination, and repair ≤2.7 × 10−2 4
Nucleic acid metabolism ≤4.6 × 10−2 5
Small molecule biochemistry ≤4.6 × 10−2 14
Physiologic system development and function
Hematologic system development and function ≤4.9 × 10−2 6
Cell-mediated immune response ≤3.6 × 10−2 5
Immune cell trafficking ≤4.9 × 10−2 3
Hematopoiesis ≤3.6 × 10−2 4
Connective tissue development and function ≤4.6 × 10−3 1

first and second weeks of INT (from 119.4 ± 14.6 to
139.6 ± 15.4) and subsequently declined during RED
(120.2 ± 11.5).
Vertical jump, snatch, and clean and jerk performances
Vertical jump performance was lower during and after
INT, by 5.9 ± 5.4% and 7.2 ± 6.5%, respectively, com-
pared with the initial week (62.7 ± 14.7 cm). It was
restored, but not enhanced above initial levels, after RED
(Fig. 6). Maximal snatch was 4.3 ± 3.7% lower than
recent best competitive performance (103.6 ± 31.9 kg)
after INT, but recovered after RED (102.3 ± 31.0 kg).
Changes in clean and jerk performance (competitive
best = 133.6 ± 40.9 kg) did not reach statistic signifi-
cance (Fig. 6).
Discussion
These data provide evidence-based insight into the
physiologic, psychologic and performance conse-
quences of a short-term, modest reduction in training
load subsequent to a period of INT training, in competi-
tive weightlifters. Key findings were that enhanced
“stress responses,” evidenced by systemic indices of cel-
lular stress, damage and inflammation, a more negative
mood state, and performance decrements, were induced
by the INT training, and largely resolved after the RED
training.
Competitive weightlifting and weightlifters are argu-
ably a relatively unique population, even among strength
and power athletes. In particular, the training is almost
exclusively high-intensity resistance exercise, and per-
formed more frequently than that prescribed for the devel-
opment of strength and power in athletes competing in
other sports (Storey & Smith, 2012). The athletes in this
study were well-trained and experienced competitors.
They were in compliance with WADA anti-doping regu-
lations under the National Sport Organization and had not
previously reported a positive test. Their mean best com-
petitive performance levels were, for snatch and clean and
jerk, 68.2 ± 7.3% and 72.5 ± 9.6% of the respective world
records at the onset of the study. Importantly, the intensity
and volume of the weightlifting training performed
during INT and RED matched that prescribed by coaches
with the intention of improving performance during the
subsequent competitive period within the annual training
program (Häkkinen et al., 1987; Fry et al., 2000; Pistilli
et al., 2008). The results of this study thus reflect the
specific nature, requirements and consequences of com-
petitive weightlifting training.
Systemic indices of cellular stress, damage
and inflammation
Gene expression in PBMCs
Our microarray data, although from a limited number of
weightlifters, add to the limited evidence from two other

34
 Stress responses in competitive weightlifters
(a) 25.0
Resting Time P < 0.05
2.0 0 h post-weightlifting
3 h post-weightlifting
Resting
3 h post-weightlifting
* 20.0
Time P < 0.05
*
CCL4 mRNA (fold change)

1.5 Time × Training load P < 0.05

CCL4 (pg/mL)
15.0 *
*
*
1.0
10.0

0.5
5.0

0.0
0.0 Intensified Reduced
Initial Intensified Reduced
Training load

(b)
Fig. 3. Plasma CCL4 concentration at rest, and 0 and 3 h after
4.0
the same 90 min weightlifting training session performed after 2
* ** weeks of intensified and 1 week of reduced training. Values are
3.5
Time P < 0.05 shown as mean ± standard deviation. Post-hoc comparisons
CXCR4 mRNA (fold change)

3.0 Training load P < 0.05 where main effects of time or training load were found: *Differ-
ent from resting level within the same training load (P < 0.05).
2.5

2.0 0.40
Resting Time P < 0.05
0 h post-weightlifting Training load P < 0.05
1.5 0.35 3 h post-weightlifting Time × Training load P < 0.05
** *
**
Protein carbonyls (nmol/mg)

1.0 0.30

0.5 0.25 *
0.0 0.20
Initial Intensified Reduced

0.15
(c)
2.0
0.10

* ** * 0.05
DDIT4 mRNA (fold change)

1.5
Time P < 0.05 0.00
Training load P < 0.05 Intensified Reduced

Training load
1.0
Fig. 4. Plasma protein carbonyl concentration at rest, and 0 and
3 h after a 90-min weightlifting session performed after 2 weeks
of intensified and 1 week of reduced training. Values are shown
0.5
as mean ± standard deviation. Post-hoc comparisons where main
effects of time or training load were found: *Different within
training load (P < 0.05). **Different at same time point between
0.0 training loads (P < 0.05).
Initial Intensified Reduced

Training load

research groups who assessed the transcriptional regula-

Fig. 2. Peripheral blood mononuclear cell mRNA expression at
rest and 3 h after the same 90 min weightlifting session per-
formed after 2 weeks of intensified training and after 1 week of
reduced training. The mRNA levels of (a) CCL4; (b) CXCR4 and
(c) DDIT4 are expressed as a fold change from resting levels
during the initial training week. Values are shown as
mean ± standard deviation. Post-hoc comparisons where main
effects of time or training load were found. *Different from
resting within the same training load (P < 0.05). **Different at
same time point between training loads (P < 0.05).
tion and responsiveness of blood cells to high-intensity
exercise in athletically trained adults (Carlson et al.,
2011; Neubauer et al., 2013). We found that after 1.5 h of
intense weightlifting in a fed state, further feeding and
3 h of recovery, PBMC showed differential expression of
genes associated with hematologic and immunologic
functions (including inflammation) and related intra- and
inter-cellular signaling and interactions. Several genes
associated with cell cycle, maintenance, including DNA
replication, recombination and repair, and death were
also affected (Table 2).

35
Storey et al.
(a)
12 80
4
DALDA part B 'worse than normal' responses
2
10

Change in performance (%)

60 0 * *

Mean (SD) per week

8
–2

6 40
–4
*
–6
4
–8
20

2 –10

–12
0 0 Vertical jump
Week 1 2 3 4 –14
Initial Intensified Intensified Reduced
Training load
Initial Intensified Intensified Reduced

Fig. 5. Daily Analysis of Life Demands of Athletes (DALDA)

(b)
Part B: Signs and Symptoms of Stress. Frequency of signs and
4
symptoms of stress reported as “worse than normal” during
initial, intensified, and reduced trainings in weightlifters. Mean 2
daily and mean (standard deviation, SD) weekly scores are *
0

Change in performance (%)

shown. *Different from intensified and reduced training weeks
(P < 0.05). –2

–4

Our analyses identified two (immune) pathways and –6

eight biologic functions in common with those from the –8

sole previous analysis of the transcriptional responses of –10
PBMC to resistance exercise (Carlson et al., 2011). Snatch
–12
Carlson et al. (2011) found differential expression
–14
of genes having immune and inflammation-related
functions, including the recruitment of leucocytes to
Best Intensified Intensified Reduced
peripheral tissues, and hematologic and cellular commu-
nication roles, 2 h after 30 min of moderate intensity (c)
resistance exercise, performed after an overnight fast, in 4

resistance trained men. 2

The activation of pro-inflammatory signaling path-
Change in performance (%)

0
ways and genes associated with the innate immune
–2
response in recognition of tissue damage has also been
–4
shown by transcriptional profiling of neutrophils 3 h
after 2 h of intense endurance exercise in fed, endurance –6

trained men (Neubauer et al., 2013). Although genes in –8

our set were also often associated with the inflammatory –10
response and other immune functions, only CXCR4,
–12
which showed a 3.8-fold increase, was in common with Clean and jerk
–14
the representative genes shown from the differentially
related gene sets found by Neubauer et al. (2013).
Best Intensified Intensified Reduced
The 1.6-fold increase in CCL4 mRNA seen in our
study with weightlifting after INT suggests a possible Training load
“priming” of the circulating mononuclear cells prior to
Fig. 6. Changes in performance relative to during initial train-
their uptake into injured tissues, including skeletal ing: (a) vertical jump, or recent maximal lifts in competition;
muscle (Fig. 2). In the earlier described study of Carlson (b) snatch; and (c) clean and jerk, with two weeks of intensified
et al. (2011), a >1.7-fold decline in CCL4 mRNA was training and one week of reduced training. Values are shown as
found after the 2 h of recovery after 30 min of resistance mean ± standard deviation. *Different from performance during
exercise. However, an earlier study, from a different initial training or previous competitive best (P < 0.05).
group, showed 2.9-fold increases in CCL4 mRNA after
30 min of cycling exercise, a 3.4-fold decline after an these data provide indications that the timing, magnitude
hour of subsequent recovery, and no difference between and rate of resolution of the transcriptional response of
pre-exercise and 1 h recovery values, in untrained men PBMC to exercise are sensitive to the mode and intensity
(Connolly et al., 2004). We suggest that, collectively, of the exercise, potentially exacerbated by high-intensity

36

eccentric muscle contractions related to tissue damage,
and the duration of the exercise. We consider our results
to reflect a response peculiar to a high-intensity, compre-
hensive weightlifting session at the conclusion of 2
weeks of INT training.
With the increases in CCL4 mRNA levels in PBMC,
we expected increases in the corresponding CCL4
protein secreted from circulating mononuclear cells
(Fig. 3). However, the CCL4 protein levels in plasma
did not differ between INT and RED, suggesting either
insufficient time for synthesis and secretion (in one or
both of INT or RED), or a low synthesis/secretion of
the protein by circulating cells or from tissues into the
circulation. The first is unlikely as no changes in plasma
CCL4 were found up to a day after resistance exercise
in recreationally strength-trained men (Risøy et al.,
2003) or muscle damage-inducing maximal eccentric
contractions of the knee extensors in non-strength-
trained men (Paulsen et al., 2005). However, we
acknowledge that the concentrations of CCL4 observed
were, for the most part, at the calculated sensitivity of
the “high-sensitivity” assay kit that we used. The same
commercial assay kit was also used in the investigations
mentioned earlier, although the specific CCL4 concen-
trations were not reported (Risøy et al., 2003; Paulsen
et al., 2005).
The combination of the increased CCL4 mRNA in
PBMC with weightlifting after INT and similar, low
levels of CCL4 in plasma after INT and RED, suggests
that there is a low synthesis/secretion of the protein by
circulating cells and, given a particular training status
and stimulus, that the chemokine is secreted from cells
only once they are within tissues and thereafter largely
retained within those tissues.
Our increased CXCR4 mRNA in the PBMC of
weightlifters 3 h after 1.5 h of training (Fig. 2) is similar
to that recently reported in neutrophils 3 h after 2 h of
high-intensity endurance exercise in trained men
(Neubauer et al., 2013). An increased CXCR4 protein
expression subsequent to the observed increase in
CXCR4 mRNA after weightlifting would draw the cells
towards, and increase their uptake into SDF-1-
expressing tissues, thereby supporting recovery from
the INT training (i.e., during RED). We propose that
CXCR4-expressing blood cells contribute to the recovery
of skeletal muscle after intense exercise in trained ath-
letes, and that this tenet should be explored in future
investigations.
The observed increases in DDIT4 mRNA in PBMC
induced by weightlifting after INT, but not RED could
reflect a summative response to a number of signals
from the organs and tissues through which the cells
have passed (Fig. 2). Because the mTOR signaling
pathway is also highly relevant in skeletal muscle, the
data suggest further exploration of the gene and its
protein in circulating cells with variations in
weightlifting training load.
Stress responses in competitive weightlifters
We can preclude an effect of diurnal variation (elapsed
time) or feeding (or lack thereof) in comparing between
the resting and post-weightlifting mRNA levels (or
changes from resting to 3 h after weightlifting) between
the INT and RED training. As the resting mRNA levels
were no different, and the same weightlifting session
was performed after both INT and RED, we propose that
the PBMCs were more susceptible or responsive to
exercise-associated stimuli after the INT training.
We have interpreted the weightlifting-induced
increases in mRNA as increases in transcript levels
within individual cells as any changes in the number or
subset proportions of PBMC would be expected to be
minor (Risøy et al., 2003; Nieman et al., 2004), and
more importantly, similar between the INT and RED.
Protein carbonyls
Our data confirm and extend prior evidence that plasma
protein carbonyls are associated with INT weightlifting
training (Margonis et al., 2007). Margonis et al. (2007)
showed that basal plasma protein carbonyl concentra-
tions were elevated following periods of INT resistance
training, inclusive of weightlifting exercises, and four
subsequent days of rest, in healthy men. In particular,
after 3 weeks of 87% INT training within the 12-week
program, and coincident with impaired maximal power
clean (−6.4%) and vertical jump (−9.6%), protein car-
bonyls were 73% higher than at the start of the program.
After a subsequent 3 weeks of RED training and 4 days
of rest, protein carbonyl levels returned to those at the
start. The resting values in our study (Fig. 4) may thus
reflect the effects of the previous training session
approximately 14 h beforehand, but similarly after both
INT and RED.
We have also provided further evidence that plasma
protein carbonyl concentration can increase during the
early recovery from high-intensity weightlifting in
trained participants (Hudson et al., 2008) and propose
that this may only be the case where the initial levels are
not already elevated. Instead, after a period of INT train-
ing, the higher resting protein carbonyl levels may
decline after weightlifting. An analysis of the particular
proteins targeted for carbonylation or decarbonylation is
of prospective interest in elaborating on the possible role
of oxidative stress after weightlifting, similar to recent
findings presented for endurance exercise (Guidi et al.,
2011).
Questionnaires
The available tools for the measurement of the sources
and symptoms of stress and mood state have not been
specifically developed nor frequently applied in com-
petitive strength and power athletes. Previously, the
DALDA has been shown to be effective in describing
differences in sources and symptoms of stress between
37
Storey et al.
training days, rest days and competition days in rugby
union players (Nicholls et al., 2009). Both DALDA
and POMS scores have shown correspondence with
periods of INT and/or RED training in endurance ath-
letes such as swimmers (Morgan et al., 1988) and
cyclists (Halson et al., 2002; Faude et al., 2009; Witard
et al., 2011).
In this study, the increases in the sources, signs and
symptoms of stress (DALDA) during the first week of
INT confirm a psychologic/affective impact, including
the perception of physiologic changes, of the doubling of
training load on the athletes (Fig. 5). The more negative
affect (global mood state) and sustained higher fre-
quency of “worse than normal” signs and symptoms of
stress during the second week of INT largely identified
the athletes as being very sore, tired. and irritable more
often. That only the global mood state, and not signs and
symptoms of stress, declined with the RED training,
suggests that there were perceivable physiologic changes
not yet completely resolved within the week. Thus,
while this study indicates that these questionnaires are
applicable in weightlifting, their meaning and effective-
ness needs further substantiation.
Vertical jump, snatch, and clean and jerk performances
The improvement in competitive performance of senior
(i.e., > 20 years old) weightlifters can be very small (e.g.,
1–2% per year; Häkkinen et al., 1987; Häkkinen et al.,
1988). The intra-athlete variation in performance in
international-level competition is also small (2.3–2.7%)
(McGuigan & Kane, 2004). Furthermore, for snatch and
clean and jerk, by definition (i.e., in competition), a
difference of 1 kg or 0.5–1.5% of the best performances
of the athletes in the current study, is meaningful. Thus,
the mean 4% lower than recent best snatch performance
after INT can be considered a robust effect (Fig. 6).
Previously, neither 1 week of similarly INT (double
load) training in junior weightlifters (Warren et al.,
1992; Fry et al., 1993, 1994; Stone & Fry, 1998) or 2
weeks of INT training in elite Finnish weightlifters
(Häkkinen et al., 1987) affected snatch or clean and jerk
performances. That snatch performance was lower after
INT, whereas clean and jerk was not, confirms a particu-
lar specificity and/or sensitivity of the lifts to the pre-
scribed training. Our finding of a 7% mean decline in
vertical jump after INT compared with the initial week
was greater than that reported previously (4%) with
increases in training load in junior weightlifters (Fry
et al., 1993). However, in a similar group of junior
weightlifters, a week of INT training also resulted in a
3% mean improvement in vertical jump height (Stone &
Fry, 1998).
That snatch and vertical jump performances recovered
to best/initial levels after a week of RED training con-
firms the expectation of an improvement in performance
(Stone & Fry, 1998; Fry et al., 2000). The magnitude of
38
improvement was comparable with that seen with INT
training followed by two or more weeks of RED training
(Häkkinen et al., 1987; Warren et al., 1992; Fry et al.,
1993, 1994). For instance, in Fry et al. (2000), 1 week of
INT training followed by 3 weeks of RED (“normal”)
training in a training camp environment, resulted in
improved (1.0–1.6%) performance in competition. Stone
and Fry (1998) reported a 3.8% mean improvement in
snatch after a week of INT (double load) training and 2
weeks of subsequently RED training, in junior
weightlifters. Similarly, small (2%) net gains in perfor-
mance at a primary competition after a 6-week cycle
consisting of 2 weeks of INT training followed by 2
weeks of normal training (−10% load) and 2 weeks of
subsequently RED (−25% load) training have been
reported (Häkkinen et al., 1987). Hence, we propose that
where a reduction in performance is incurred during INT
training, a week of RED training is very likely to enable
the restoration of performance, whereas a further 1–3
weeks of RED training may result in improvement above
the earlier initial levels.
The necessity and value (benefit or harm) of the INT
training to improving competitive performance cannot
be confirmed from the current data. However, the reso-
lution of the stress responses and the performance dec-
rement (incurred during INT) before or by the end of
RED (lower volume, but sustained high intensity) argues
against the training being detrimental or debilitating.
That the performance of the weightlifters in this study
was restored after the combined INT and RED training is
consistent with accepted paradigms and terminology
from applied sport science literature that the training
represented an “overload” resulting in “overreaching,”
and not “overtraining” (Meeusen et al., 2013). Thus, the
INT training may be necessary/effective/preferable for
(subsequent) improvements in performance in experi-
enced competitive weightlifters. Further evidence, albeit
problematic to obtain with competitive athletes, is
required to substantiate this supposition.
These results implicate CXCR4-expressing monocytes
and chemokine signaling in the early recovery from
weightlifting at the end of a period of INT training.
Coincident with increased resting plasma protein car-
bonyl concentration, negative affective responses and
decrements in performance, the changes in CXCR4,
CCL4, and DDIT4 mRNA expression in PBMCs may
represent an exacerbated response to early signals from
injured or inflamed tissues, including skeletal muscle.
The absence of this response to the same training per-
formed after a subsequent week of RED training sug-
gests a resolution of peripheral and/or intracellular
signals associated with the INT training.
Conclusion
This study has identified systemic changes, as contribu-
tors within an apparent comprehensive stress response

system, to the recovery from intense weightlifting and
changes in performance with variations in training load.
Two weeks of INT training resulted in systemic indices
of cellular stress, damage and inflammation, increases in
psychologic stress, and reduced snatch and vertical jump
performances. A subsequent week of RED training was
sufficient to resolve most changes, and restore, but not
enhance performance.
Perspectives
This study adds to the sparse evidence of the conse-
quences of short-term variations in training load in
competitive weightlifters. The findings implicate
chemokine signaling in the stress response to intense
weightlifting, particularly a role for circulating
CXCR4-expressing mononuclear cells in the recovery
of skeletal muscle that warrants further investigation.
Lastly, the evidence suggests that short-term reductions
Stress responses in competitive weightlifters
in training load may be important in ensuring tolerance
to, and recovery from, periodic INT training in strength
and power athletes.
Key words: Snatch, clean and jerk, overload, recovery,
peripheral blood mononuclear cells, cDNA microarray,
gene expression, psychologic status.
Acknowledgements
Funding for this work was provided by the University of
Auckland Postgraduate Research Student Support scheme
(A. G. S.) and Faculty of Science (H. K. S.). We thank Liam
Williams for providing expert technical advice in performing
the microarray experiments in the Centre for Genomics,
Proteomics, and Metabolomics. P. C. Tong and Stefan Wette are
also thanked for their assistance with the qRT-PCR. NorthSport
Olympic Weightlifting and AUT Millenium Institute of Sport and
Health generously granted us use of their facilities. Lastly, the
athletes are thanked for their efforts and commitment to the
study.

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