A systematic review of potential productivity, egg quality, and animal welfare

implications of extended lay cycles in commercial laying hens in Canada

Vivek Arulnathan ,*,1 Ian Turner ,* Nicole Bamber ,* Jannatul Ferdous ,* Florian Grassauer,*
Maurice Doyon ,y and Nathan Pelletier*
*
FIP 226, Food Systems PRISM Lab, Fipke Centre for Innovative Research, University of British Columbia
Okanagan, Kelowna, BC, V1V 1V7, Canada; and yPaul-Comtois, Laval University, Quebec City, QC, G1V 0A6,
Canada

ABSTRACT Lay cycle lengths in the Canadian egg
industry are currently 50 to 52 wk (68−70 wk of age). In
light of increased productivity in commercial laying hens
over the last few decades, the much longer lay cycle lengths
already implemented in other countries, extending lay
cycle lengths in Canada, should be considered with careful
attention to potential environmental, economic, and ani-
mal welfare implications. However, there is a lack of infor-
mation in the public domain that provides robust evidence
of performance levels and potential trade-offs to support
comprehensive consideration of the desirability of extend-
ing lay cycles beyond current Canadian norms. Hence, a
systematic literature review using the Preferred Reporting
Items for Systematic Reviews and Meta-Analyses
(PRISMA) methodology was undertaken. Data collec-
tion focused primarily on information related to feed-,
flock management-, and hen genetics/physiology-related
interventions that were studied in literature to support
extension of lay cycles (review objective 1), and compiling
and analyzing productivity, egg quality, and animal
Key words: egg production, extended lay cycle, productivity, egg quality, animal welfare
https://doi.org/10.1016/j.psj.2024.103475
welfare outcomes reported at 70 wk of age or beyond
(review objective 2). Several feed-related interventions
such as high-protein diets, and probiotics supplements,
and flock management interventions such split-feeding
were found to potentially improve productivity, and espe-
cially egg quality, outcomes in the late laying phase. More
studies with bigger flock sizes and in commercial lay facili-
ties need to be undertaken before any of these interven-
tions can be definitively recommended for commercial egg
production. Under objective 2, productivity was found to
be at acceptable levels well beyond 70 wk of age. Perfor-
mance on most egg quality traits and animal welfare indi-
cators were also at acceptable levels past 70 wk of age but
increased variability was observed beyond »80 wk of age.
There were also inconclusive indications on how hens in
caged housing and white laying hens fare relative to hens
in noncaged housing and brown-type layers during the
late laying phase. Economic data were limited but sug-
gested that lay cycle lengths beyond 90 wk might not gen-
erated net economic benefits.
2024 Poultry Science 103:103475

INTRODUCTION practice is the extension of lay cycles beyond the norms

The modern Canadian egg industry achieves consider-
ably improved productivity, resource efficiency, and ani-
mal health outcomes relative to historical performance
levels under current management practices (Pelletier,
2018). However, there remains significant potential for
continued improvement through identifying and dissem-
inating sustainability best management practices (Pel-
letier, 2017). One potentially promising management
Ó 2024 The Authors. Published by Elsevier Inc. on behalf of Poultry
Science Association Inc. This is an open access article under the CC
BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/
4.0/).
Received November 10, 2023.
Accepted January 12, 2024.
1
Corresponding author: vivek.arulnathan@ubc.ca
currently practiced in Canada. Depopulation of flocks in
Canada is normally done between 68 and 70 wk of age
(Turner et al., 2022) (i.e., after 50−52 wk of lay).
Extending lay periods beyond this, as is already common
elsewhere in the world, may potentially have environ-
mental, animal welfare, and economic implications, but
the potential synergies, benefits, impacts, and/or trade-
offs within and across these areas of concern are unclear.
Historically, lay cycles in commercial egg production
have been extended through induced molting, which is a
controversial practice (NFACC, 2017). This process is
associated with increased productivity and improved
egg quality compared to at the end of pre-molt cycle,
reduced costs, and reduced investments in breeders and
hatcheries (Yousaf and Chaudhry, 2008). Molting by
feed withdrawal is considered the most effective practice

1
2 ARULNATHAN ET AL.
to extend the productivity of hens, but is also associated
with welfare concerns such as hunger, loss of immunity,
increased mortality, and increased contamination of
eggs (Holt, 2003; Yousaf and Chaudhry, 2008). The
practice of induced molting by feed withdrawal is
banned in Europe and is not recommended except in
emergencies in Canada. However, keeping hens in lay up
to 100 wk of age without molting may nonetheless be
realized if lay persistency and maintenance of egg qual-
ity can be achieved by other means (Bain et al., 2016).
The global egg industry has seen consistent increases
in hen productivity over the last few decades. Between
1990 and 2010 alone, hens increased average production
by »2.5 eggs every year. In addition to lay cycles extend-
ing from 68 to 75 wk of age on average, this has led to an
increase in rate-of-lay of 70 eggs/hen/lay cycle in 2 deca-
des (Sambeek, 2011a). Canada’s relatively short lay
cycles may mean that hens are depopulated despite still
having acceptable levels of lay persistence and egg qual-
ity. Data obtained from Hendrix Genetics (personal
communication) show, for example, that average lay
cycles in the Netherlands are now roughly 90 wk long for
white hens (cage), and 80 wk long for brown hens (avi-
ary). Further, breeding companies have stated that
achieving commercial layers capable of producing “500
eggs in a single production cycle of up to 100 wk of age”
is a realizable objective in commercial egg production
(Sambeek, 2011b; Anco, 2018).
However, several factors are critical to successfully
extending lay cycles. Currently, the main reason for
replacing laying hens is deteriorating egg quality in the
later stages of the lay cycle (Fathi et al., 2019). Dunn
(2013) suggests that the decrease in egg quality and lay
persistency does not mean that all hens produce poor-
quality eggs but rather indicates increased variability in
egg quality in a flock. This variability in lay persistency
is attributed to the subtle differences in the neuroendo-
crine system of birds. Thus, maintaining the tissues and
organs responsible for egg production is considered the
most important prerequisite for extending lay cycles
(Bain et al., 2016).
Given Canada’s relatively shorter lay cycle lengths,
extending lay cycles can potentially be a strategy with
significant environmental, economic, and animal welfare
implications. However, there is a lack of robust and reli-
able information in the public domain on the performance
levels achievable by commercial layers in extended lay
cycles. To the best knowledge of the authors, there have
been no efforts yet to systematically review and analyze
the scientific literature on extended lay cycles. To provide
evidence of the potential sustainability implications of
extending lay cycles to egg producers, it is important to
first determine the performance levels achievable by lay-
ing hens beyond 70 wk of age (average age of depopula-
tion in Canada). Given the unique challenges associated
with extending lay cycles, it is also important to identify
the strategies and practices that will enable the mainte-
nance of lay persistency and egg quality without
adversely impacting welfare outcomes. Hence the objec-
tives of the current review are to:
1. Identify management practices, strategies, and tech-
nologies that may enable the successful extension of
lay cycles beyond »50 to 52 wk lay period.
2. Compile and analyze information on the potential
productivity, egg quality, and animal welfare implica-
tions of extending lay cycles for laying hens beyond
70 wk of age.
This systematic literature review will be beneficial to
several stakeholders. It will help in screening and identi-
fying the performance levels and strategies for Canadian
egg farmers to consider for extending lay cycles. It will
also provide researchers with a comprehensive summary
of the current state of research and help in identifying
literature gaps with respect to extended lay cycles.
METHODS
PRISMA Methodology
To identify literature addressing the extension of lay
cycles, a systematic review of poultry science literature
was undertaken using the Preferred Reporting Items for
Systematic Reviews and Meta-Analyses (PRISMA)
methodology. The PRISMA method is a set of evidence-
based, minimum requirements for systematic reviews
that enables authors to transparently report the why,
how, and what of literature reviews (Page et al., 2021).
Search Protocol The search parameter “(ALL=
(“lay*”) AND ALL=(“egg*”) AND ALL=(“extend*”))”
was used to search for literature in the Web of Science
(WoS) Core Collection database in February 2022. The
following criteria were used to identify papers for inclu-
sion in the review.
 Only literature published in the last decade
(2011−2021) was considered to ensure relevance to
current practices in egg production.
 Only literature in English was considered.
 Review papers and conference proceedings were not
considered for data extraction.
The search results were imported into the Covidence
screening and data extraction tool (Covidence, 2022).
The titles and abstracts were initially screened to deter-
mine if papers specifically mention extended lay cycles
or discuss any aspects of hen productivity, egg quality,
animal welfare, or the economics of egg production that
might be relevant for extended lay cycles. This screening
process was carried out by 2 of the authors indepen-
dently in Covidence to minimize the risks associated
with biased selection of papers for review. Conflicting
decisions by the two screening authors were resolved by
discussing the abstracts together and reaching a consen-
sus decision.
Once the initial shortlist was finalized, the papers
were then screened to identify the lay cycle length stud-
ied. Papers were selected for data extraction only if they
addressed one or more of productivity, egg quality, eco-
nomics, or animal welfare considerations associated with
 EXTENDED LAY CYCLES LITERATURE REVIEW
hens older than 70 wk of age (50−52 wk of lay). Any
papers not meeting this criterion were excluded and not
considered further. This second round of screening was
done by only one author since a clear objective criterion
was defined for inclusion.
In addition to the search of literature using the WoS
database, a secondary search was done using Google
Scholar. This was necessary since research has shown
that relevant literature can be missed if only one search
engine/database is used in a systematic review (Hadd-
away et al., 2015). The same search parameters used in
WoS were employed in Google Scholar. The same pro-
cess for title and abstract screening and shortlisting for
data extraction described above was employed for the
search results. The title and abstract screening in Google
Scholar was limited to the first 200 results as recom-
mended by Haddaway et al. (2015), since the relevance
of the results beyond this point is often questionable.
The shortlist generated was compared against the short-
list from WoS to remove any duplicates.
General Information Once the final list of papers for
data extraction was determined, general information to
characterize each study was extracted. Data extracted
under this category included the following: year of study,
the type (s) of housing system, bird type(s), location, and
lay cycle length. The housing system used was classified
as one of conventional, enriched, aviary/free-run, or free-
range systems. For cage-type housing, information on
whether the hens were housed individually for the study
was also documented. For classifying the bird type, the
color of the hens (“white” or “brown”, and the genetic
strain (e.g., Bovan) was used. The country in which the
study was conducted was used to characterize the loca-
tion of the study. The age of hens when placed under
study conditions and the age of hens at the end of the
study were documented to determine the lay cycle length
studied. Any reported preparatory/nonobservation peri-
ods under study conditions at the beginning were not
considered in the calculation of lay cycle length.
Review Objective 1
The purpose of the first objective was to identify and
describe potential strategies, practices, and technologies
that can support keeping hens in lay beyond the age of
70 wk. For clarity, all strategies, practices, and technolo-
gies considered are referred to as “interventions.” For this
objective, the data were extracted under 2 categories:
1) interventions studied and 2) study conditions. Under
the “interventions studied” category, the following infor-
mation was collected: name of the intervention, cate-
gory, description of the intervention, intended effect,
and scenarios studied. An initial list of categories includ-
ing feed ingredients, flock management, animal welfare,
environmental conditions, and genetics was defined to
classify interventions. The intended effect described the
hypothesis stated by the authors regarding an interven-
tion. For example, Moln
ar et al. (2018) hypothesized
that providing hens with different diets during different
3
times of the day (split feedings) would potentially
improve calcium availability in older laying hens and
thus improve egg quality.
Additionally, data were extracted in objective 1 to
characterize the conditions under which the study was
conducted. This helps in contextualizing both the inter-
ventions (identified under this objective) and the perfor-
mance of hens (documented in review objective 2).
Information collected under the “study conditions” cate-
gory included: flock size; number of lay periods studied,
lighting regime, environmental conditions (temperature
and humidity), feed composition, and feed and water
availability. Whether or not induced molting was used
was recorded where relevant. Even though the focus of
this review was on single-extended lay cycles, the few
studies related to molted hens shortlisted were not
excluded as they either supported comparison with non-
molted hens (Regmi et al., 2018) or addressed issues
such as bone strength and hen physiology (Wein et al.,
2020) that could be relevant for old hens in extended
production cycles. For feed composition, the percent
content of ingredients in the base diet and any changes
to specific ingredients in the different scenarios were
recorded.
Review Objective 2
This review objective aimed to determine the produc-
tivity, egg quality, animal welfare, and economic out-
comes achievable in lay cycles beyond 70 wk of age. The
data extracted here included: indicators used to measure
performance outcomes, indicator category, age of birds
when results were recorded, and indicator results. Indi-
cators were categorized into one of the following: pro-
ductivity, egg quality, animal welfare, and economics.
Indicator results were reported both at a specific age (e.
g., tibia strength at 100 wk of age) or over a monitoring
period (e.g., mortality rate between 84 and 95 wk of age)
in the literature reviewed. Results for all the indicators
used to analyze performance in a study were collected.
Results of baseline performance were also recorded, if
available, for comparison.
For each shortlisted paper, data extraction was per-
formed by only 1 author. Unlike the initial shortlisting
of papers where a reviewer had to make a subjective call
on a paper’s inclusion, the data extraction process had
minimal risk of bias since no subjective information was
being collected. If any required information was not
explicitly available in a paper, no assumptions or substi-
tutions were made.
Once data extraction for review objective 2 was com-
plete, commonly used indicators in the literature
reviewed for each category were identified. Only indica-
tors for which data were available from 3 or more publi-
cations reviewed are presented in the “Results and
Discussion” section. Finally, the performance of laying
hens for these commonly used metrics was then analyzed
relative to age, and if possible, relative to housing system
type and hen breed.
4 ARULNATHAN ET AL.
RESULTS AND DISCUSSION
Literature Identification
The initial search of literature in the WoS database
produced 544 results. Fifty-six of these papers were sub-
sequently shortlisted for inclusion, from which 20 were
selected for data extraction.
Employing the same search criteria in Google Scholar,
an additional 11 papers were selected for data extrac-
tion. One of these papers (Flock, 2016) was analyzing
the results from multiple North Carolina layer perfor-
mance and management tests and, hence, the reports of
these four tests were obtained and added to the list for
data extraction (with Flock (2016) excluded). In total,
34 unique publications were selected for data extraction.
Figure 1 provides a visual representation of the
PRISMA-based literature screening and identification
process and results as described.
General Information
At least one publication from each of the years
between 2011 and 2021 was included in the final list for
data extraction—except 2013 and 2014. The growing
interest in extended lay cycles and the proliferation of
literature in this domain is evidenced by the fact that 23
of the 34 papers identified for review were published in
2017 or later.
Studies addressing lay cycles in conventional cage
layer housing were most common—with 25 of the 34
publications in the final list studying extended lay cycles
in conventional housing systems either exclusively or
alongside other housing systems. Enriched (colony)
Figure 1. Representation of the literature identification process using the PRISMA flow diagram from Page et al. (2021).
housing was included in 8 publications and free run/avi-
ary housing systems in 11 of the 34 publications. Free-
range housing was the least represented housing system
(3 publications). Eight of the 34 publications in the final
list studied more than one housing system, with Ander-
son (2018) and Anderson (2015) both including 4 differ-
ent housing systems.
Reporting on the housing systems used at the pullet
phase was found in only 7 of the 34 publications. This is
not surprising given that most studies acquired and
housed the birds under experimental conditions exclu-
sively for the late-laying phase. Among the studies that
reported the type of pullet housing, conventional caged
pullet rearing was the most common.
A total of 41 different breeds of laying hens were ana-
lyzed in at least one of the publications reviewed—with
both brown (16) and white (25) type laying hens well-
represented. Over 50% of these layer types (23 of 41)
were analyzed exclusively in 1 of the 4 North Carolina
layer performance tests included in the review
(R23−R26 in Table 1). Hy-Line Brown and ISA Brown
were the 2 most widely analyzed layers—each being
included in 9 of the 34 publications reviewed. Among
white type layers, Bovan White (7) and Hy-Line W36
(6) were the most studied.
Analysis of extended lay cycles from 18 different coun-
tries was considered in this review. Five publications
from each of Belgium, China, and the United States
accounted for almost half of the publications reviewed.
Experimental/monitoring periods in the publications
reviewed varied widely—from as few as 2 wk of lay (van
de Reep et al., 2018; Wein et al., 2020) to analysis of
complete 99 wk-long laying periods (Regmi et al., 2018).
The age of hens at the end of the study ranged between
 EXTENDED LAY CYCLES LITERATURE REVIEW 5
Table 1. General information related to publications from which data were extracted in this review.

Reference Age at beginning Age at end

number Reference Type of housing—layers Geography of study (wk) of study (wk)
R1 Anwar et al. (2015) Conventional Pakistan 70 77
R2 Anwar and Rahman Conventional Pakistan 71 95
(2016a)
R3 van de Reep et al. (2018) Aviary Netherlands 84 86
R4 Morales et al. (2018) Conventional Columbia NA NA
R5 Hao et al. (2021) Conventional—individually China 70 75
housed
R6 Yalcin et al. (2020) Conventional, Enriched 92 16
R7 Wein et al. (2020) Conventional Israel 73 75
R8 Abo Ghanima et al. (2020) Conventional, free-run Egypt 28 76
R9 Wall et al. (2022) Enriched, free-run Sweden 15 100
R10 Hanlon et al. (2021) Conventional Canada 1 100
R11 Sepehr et al. (2021) Conventional Iran 75 84
R12 Moln
ar et al. (2017) Enriched Belgium 72 83
R13 Guo et al. (2021) Conventional China 87 97
R14 Saibaba et al. (2021) Conventional Israel 15 86
R15 Sun and Kim (2020) Conventional—individually South Korea 63 73
housed
R16 Anwar and Rahman Conventional Pakistan 70 90
(2016b)
R17 Kara et al. (2016) Conventional Turkey 80 92
R18 Moln
ar et al. (2016) Enriched, aviary Belgium 61 83
R19 Moln
ar et al. (2018a) Aviary Belgium 17 85
R20 Alfonso-Carrillo et al. Conventional—individually Spain 98 105
(2021) housed
R21 Regmi et al. (2018) Aviary USA 17 116
R22 Sirri et al. (2018) Enriched Italy 28 81
R23 Anderson (2012a) Conventional USA 17 109
R24 Anderson (2012b) Conventional, free-run, free USA 17 85
range
R25 Anderson (2015) Conventional, enriched, USA 17 89
free-run, free range
R26 Anderson (2018) Conventional, enriched, USA 17 89
free-run, free range

R27 Tumov
a and Gous (2012) Conventional SA 83 99
R28 Liu et al. (2018) Conventional—individually China 72 80
housed
R29 Moln

ar et al. (2018b) Conventional—individually Belgium 75 92
housed
R30 Çatl{ et al. (2012) Conventional Turkey 85 105
R31 Zhang et al. (2019) Conventional China 72 78
R32 Moln
ar et al. (2014) Aviary, enriched Belgium NA NA
R33 Calik (2017) Aviary Poland 21 99
R34 Zhu et al. (2022) Conventional China 67 76

73 (Sun and Kim, 2020) and 116 wk of age (Regmi et al.,
2018). The average age of hens at the end of the laying
cycle in the publications reviewed was 89 wk. Approxi-
mately half of the publications reviewed (16 of 34) con-
ducted their analysis exclusively in the late laying phase
(70 wk of age or higher). Table 1 details some of the gen-
eral information extracted from each of the 34 publica-
tions in the final list. An expanded version of Table 1
can be found in supplementary file—Table S1.
Review Objective 1—Interventions
Interventions were categorized under 3 categories:
feed composition, flock management, and genetics/phys-
iology (physiology here refers to all body systems). Four-
teen of the 34 publications reviewed studied the effects
of different feed supplements on the performance of lay-
ing hens. The feed-related interventions investigated
were quite diverse in both the source of ingredients and
intended effects. Studies on the effects of calcium supple-
ments (Moln
ar et al., 2017), meat and bone meal
(MBM), and oyster shell meal (OSM) (Çatl{ et al.,
2012) targeted improving otherwise deteriorating egg-
shell quality in aged laying hens. Studies focused on rose-
mary and cinnamon oils (Abo Ghanima et al., 2020),
vitamin A and vitamin K3 supplements (Guo et al.,
2021), and dried pomace supplements (Kara et al.,
2016) all investigated their oxidative stability in older
laying hens. The effects of high-protein diets and diets
with probiotic and symbiotic supplements on the immu-
nity and hen performance were also investigated (Anwar
et al., 2015; Anwar and Rahman, 2016a,b). Other focus
areas for feed-related studies were the effects of supple-
ments on intestinal functions and the health and immu-
nity of hens (Zhang et al., 2019; Zhu et al., 2022). All of
these 14 studies analyzed performance in extended lay
cycles using productivity and egg quality metrics, but
only roughly half of the feed-related studies analyzed
animal welfare concerns.
6 ARULNATHAN ET AL.
Seventeen publications studied the effects of hen
genetics or hen physiology in extended lay cycles. How-
ever, in most of the cases of hen genetics or physiology,
the studies analyzed 1 of the following 3 aspects: 1) com-
paring different layer breeds (Hanlon et al., 2021),
2) comparing performance at different points of the
extended lay cycle (Calik, 2017), or 3) physiological
changes in layers at old age (Hao et al., 2021). Thus,
unlike the feed-related interventions, publications in the
review dealing with hen genetics and physiology rarely
addressed any specific practices or strategies that
affected performance in aged laying hens. As a result,
the term “intervention” should be used cautiously when
addressing layer genetics or physiology in the current
analysis.
Ten of the 34 publications analyzed interventions
related to flock management. Interventions categorized
under flock management were quite diverse. This cate-
gory included studies on the effects of alternative feeding
regimes (Moln
ar et al., 2018a; Saibaba et al., 2021),

ambient temperature (Tumov
a and Gous, 2012), litter
substrates, and molting (Regmi et al., 2018). The most
commonly addressed aspect under flock management,
however, was the effect of housing type (Anderson,
2012a, 2015, 2018; Abo Ghanima et al., 2020). Table S2
in the supplementary file provides a detailed summary
(name of intervention, description, scenarios, and
hypothesis) of all the interventions related to extended
lay cycles identified in the reviewed literature but some
of the key findings reported in these studies are discussed
below.
Feed-related interventions identified in this review
produced a positive impact in many studies on the pro-
ductivity, egg quality, and welfare outcomes in extended
lay cycles—though further exploration of some of the
highly promising feed supplements would be required
before recommendation for use in commercial egg pro-
duction. High-protein diets (18% crude protein [CP])
and diets with probiotic and symbiotic supplements
(16% CP) were found to be effective in increasing the
immunity levels of 77-wk-old postmolt laying hens
(Anwar et al., 2015). This increase in immune response
in hens resulted in lower mortality rates for protein/
symbiotic/probiotic supplemented diets (2−3.5%) com-
pared to a control diet (16% CP) with no supplements
(5.5%) (Anwar et al., 2015).
A study on the productive and egg quality perfor-
mance of hens fed with similar control (16% CP, no sup-
plements) and supplemental diets found that probiotic
and symbiotic supplements have the potential to
increase egg production in older hens observed up to 90
wk of age. High-protein (18% CP, no supplements), pro-
biotic-, and symbiotic-supplemented diets (16% CP)
also increased eggshell thickness and eggshell breaking
strength, and reduced bacterial contamination of eggs,
but produced a decrease in the internal quality of eggs
(Anwar and Rahman, 2016b). Considering the positive
effects of a high-protein diet and probiotic and symbiotic
supplemented diets on older, postmolt layers, future
research that studies the performance and welfare
outcomes of using these supplements on nonmolted hens
is warranted.
The effectiveness of probiotics—in the form of Entero-
coccus faecalis supplement (at 3.75E8 cfu EF/kg and
7.5E8 cfu EF/kg inclusion rates)—was also explored in
Zhang et al. (2019). Enterococcus faecalis supplements
improved egg production and feed conversion ratio
(FCR) but did not improve egg quality. Zhu et al.
(2022) similarly explored the effect of dietary supple-
ments on gut microbiota. Adding Broussonetia papyri-
fera fermented feed (best results seen at 3% inclusion
rate, 15.6% CP)—which contains antioxidant and anti-
inflammatory properties—as a replacement for soybean
meal (control diet, 15.72% CP) improved the diversity
of gut microbiota and showed signs of potentially
improving the intestinal health of hens (Zhu et al.,
2022).
Rosemary and cinnamon essential oils (included at
300 mg/kg in corn-based control diet with 18% CP,
2.81% crude fiber, 3.85% calcium, and 3,800 kcal/kg
metabolic energy)—considered effective due to their
antibacterial, antioxidative, and anti-inflammatory
properties—produced an increase in egg production
(63% laying rate in groups fed oil supplemented diets vs.
56% in control group) and better FCR (3.1 vs. 3.6 kg of
feed/kg of eggs) during 68 to 76 wk of age. The study
reported better performance throughout the laying cycle
when essential oils were included and given the potential
for essential oils to also positively impact blood chemis-
try, immunity, and antioxidant parameters, the paper
recommended a supplementation rate of 300 mL/kg of
layer feed (Abo Ghanima et al., 2020). It is important to
take into consideration that these are results from just
one study and whether similar improvements can be rep-
licated with other essential oils and also obtained in a
flock with higher baseline productivity levels needs to be
analyzed. Another plant-based by-product known for its
antioxidative properties—grape pomace supplements—
was found to have no significant impact on productivity
and egg quality (Kara et al., 2016).
Supplementing diets with organic, bioavailable zinc,
as compared to zinc made available through inorganic
mineral salts (ZnO), did not significantly affect the skel-
etal integrity (measured using keel bone deformation)
and mortality between 15 and 100 wk of age (Wall et al.,
2022). Bioavailability of essential compounds was also
the focus in studying the effects of MBM and OSM. Par-
tially replacing dietary limestone with MBM (4% inclu-
sion level), OSM (2%), or MBM + OSM (4%
MBM + 2% OSM) supplements in aged laying hens
showed promise. Oyster shell meal and MBM + OSM
supplements improved productivity of hens, whereas
MBM was found to be the most effective in improving
both external and internal egg quality (Çatl{ et al.,
2012). Moln
ar et al. (2017) and Moln
ar et al. (2018b)
also focused on the possibility of improving calcium
availability for egg formation by trialing different ratios
of fine and coarse limestone in conventional and split
feeding systems. While feeding coarse limestone enables
better retention, and flocks under split feeding had
 EXTENDED LAY CYCLES LITERATURE REVIEW
higher calcium intake, it was found that the decreased
solubility of coarse limestone supplied exclusively in the
afternoon limits the calcium available for egg formation
during dark hours. Split feeding with a 70% coarse and
30% fine limestone ratio in the afternoon was found to
be the best diet in terms of both egg quality and skeletal
integrity in this study but split feeding with limestone
made available both in the morning and afternoon diets
could potentially be more efficacious in improving the
performance of laying hens beyond 70 wk of age (Moln
ar
et al., 2017).
Deficient, adequate, and excess vitamin A (0, 7,000,
and 14,000 IU/kg) and vitamin K3 (0, 2, and 4 mg/kg)
supplements did not significantly impact on egg laying
rate, egg weight, FCR, or skeletal integrity (measured
using tibia breaking strength). On the other hand, feed-
ing adequate amounts of vitamin A (7,000 IU/kg) and
vitamin K3 (2 mg/kg) supplements was recommended
by the authors since both were found to improve eggshell
ratio, eggshell breaking strength, and the antioxidative
functions of eggshell glands even at 97 wk of age com-
pared to a control diet (1,320 IU/kg of vitamin A and
0.5 mg/kg of vitamin K3) (Guo et al., 2021).
Adding flaxseed—which is rich in omega 3 fatty acids
that influence several biological functions of hens—to
layer diets and the impact of different processing meth-
ods on productivity, egg quality, blood profile, antioxi-
dant stability, and immunity was analyzed in Sepehr et
al. (2021). Addition of 9% extruded flaxseed increased
egg laying rate and improved both internal and external
egg quality in late laying hens (75−84 wk of age) com-
pared to a control diet. Similarly, supplementing corn
and soybean-based layer diets with high microbial phy-
tase (basal diet with 0% phytase compared to 0.06% and
0.12% phytase inclusion rates) also produced higher pro-
ductivity and better egg quality, but the impact of phy-
tase on hens older than 73 wk of age was not explored
(Sun and Kim, 2020). For all feed-related interventions
discussed, refer to Table S2 in the supplementary file for
more details.
Time-restricted feeding was found to have the poten-
tial to delay the decline in performance typically seen in
aging (86 wk) laying hens. Although only one time-
restricted feeding configuration was trialed in Saibaba et
al. (2021), the results suggested further potential for
improvements in productivity and egg quality in older
laying hens with an optimized time-restricted feed regi-
men. On the other hand, analysis of split feeding systems
was affected by the flock experiencing severe health and
sustained feather pecking issues in other studies (Moln
ar
et al., 2018a,b). Nonetheless, in light of positive (if
inconclusive) outcomes with respect to eggshell weight,
eggshell breaking strength, and tibia breaking strength,
it appears that split feeding, too, has the potential to
improve the performance of older laying hens (observed
up to 85 wk of age) and hence merits further exploration
and optimization.
Increased ambient temperature (28°C vs. 20°C) was
found to have no significant impact on the performance
of 83- to 99-wk-old laying hens, though a small reduction
7
in productivity was seen due to reduced feed intake at

higher temperature (Tumov
a and Gous, 2012). Simi-
larly, litter substrates were not found to have any
major effects on the productivity and welfare outcomes
of 116-wk-old laying hens other than increased feather
loss when AstroTurf is used (Regmi et al., 2018). The
same study also compared extending lay cycles with and
without implementing a molt. Molted hens had higher
productivity and lower mortality compared to the non-
molted flock but experienced greater welfare issues such
as increased incidence of comb wounds due to aggressive
behavior (Regmi et al., 2018). Anderson (2012a) also
compared the performance of molted and nonmolted
hens and found that molted hens have better productive
and egg quality performance in the late laying phase (73
−109 wk of age). It must be noted that both of these
studies considered the performance of nonmolted hens
well past 100 wk of age, where a drastic drop in produc-
tivity and quality in single-extended cycles would be
expected (Anderson, 2012b).
Genetic selection to support 100 wk of lay has also
produced physiological effects. van de Reep et al. (2018)
found that Salmonella vaccination done at the pullet
stage could likely become ineffective late into the
extended laying period, resulting in both animal welfare
and food safety concerns. One potentially major physio-
logical change in modern commercial layers is the shift
in the trigger for sexual maturation. It is an established
fact that sexual maturation in hens requires a photoperi-
odic trigger and activation of the hypothalamic pituitary
gonadal axis. However, intensive genetic selection for
productivity has made body weight and reaching specific
metabolic thresholds become the potential trigger for
onset of lay—which enables earlier and longer peak pro-
duction (Hanlon et al., 2021). These changes in pituitary
sensitivity in turn enable repeated follicular growth,
which facilitates the maintenance of lay persistency in
modern laying hens. Suppression of this follicular devel-
opment and increased ovarian oxidative stress were
found to be influential in the decline in productivity of
aged laying hens (Hao et al., 2021). A genome-wide asso-
ciation study also revealed hen physiological aspects and
specific genes (RHOA, SDF4, and TNFRSF4) to be
associated with the maintenance of egg quality in the
late laying phase (Liu et al., 2018).
A potentially key requirement for achieving lay persis-
tency and egg quality in longer lay cycles is nutrition
and feeding practices, especially in the pullet phase. As
discussed later in section 3.3.3 with respect to building
up the medullary bone, energy/protein ratio between
ages 11 and 16 wk for pullets, adjusting feed intake dur-
ing onset of egg production, maintaining body weight,
managing decreasing energy requirements after egg pro-
duction is established, optimizing protein and amino
acid levels in the diet, adjusting calcium requirements
for different stages of the lay cycle, and feeding system/
technique used are all important in achieving longer lay
cycles (Bain et al., 2016; Dekalb, 2017; Moln
ar, 2017).
That publications addressing such pullet-related inter-
ventions were not identified in the literature reviewed is
8 ARULNATHAN ET AL.
one of the main limitations of this study—possibly due
to the combination of search terms used for identifying
literature.
Study conditions documented were largely homoge-
nous and reflected common practices such as ad-libitum
feed and water intake, and relatively consistent ambient
temperatures and lighting regimes. The median flock
size in the literature reviewed was 228 birds. Though
there were data reported from commercial flocks
(Moln
ar et al., 2014) and birds acquired after depopula-
tion from commercial barns, the literature reviewed
mostly dealt with small experimental flock sizes. Only 8
of the 34 publications reviewed had a flock size larger
than 1,000 birds. Eight of the 34 publications reviewed
dealt with molted flocks.
Fifteen of 19 publications that provided information
on lighting regimes specified 16 h of light and 8 h of dark-
ness per day. No studies focusing on the effects of alter-
nate lighting programs were shortlisted in this review,
though it has been mentioned as potentially being a fac-
tor in the efficacy of extended lay cycles (Moln
ar, 2017).
The environmental temperature ranged between 20 and

23°C. Only 1 paper (Tumov
a and Gous, 2012) specifi-
cally studied the impacts of ambient temperature on
older laying hens. Humidity was documented in less
than 15% of the reviewed literature. Feed compositions
specified were predominantly corn-based, with soybean
and wheat being other major ingredients. Provision of
limestone in the late laying phase ranged between 7 and
10% of the feed mass across all of the studies. Most of
the studies referred to the NRC requirements for layer
nutrition (NRC, 1994) as the basis on which the energy
and nutritional requirements were determined. None of
the publications reviewed documented water consump-
tion. Overall, there was insufficient variability in the
study conditions to enable meaningfully differentiating
or contextualizing the influence of these variables on the
productivity, egg quality, or animal welfare outcomes
discussed in the results of review objective 2. The table
summarizing study conditions observed in the
literature reviewed is provided in the supplementary file
(Tables S3 and S4).
Review Objective 2
This section presents the performance of hens in
extended lay cycles for a subset of productivity, egg
quality, animal welfare, and economic indicators. The
indicators presented and discussed here are those that
were commonly reported in the literature reviewed.
Productivity Productivity of laying hens was primarily
reported in terms of egg laying rate (%) and average egg
mass (g). Egg laying rate was mostly measured on a hen
daily basis. The average reported laying rate in lay
cycles beyond 70 wk of age was 74% and the median was
76%. If only the best reported laying rate from each
study is considered, the mean and median increased to
76 and 79%, respectively. The highest reported laying
rates beyond 70 wk of age were »89% between 68 and
76 wk of age (Zhu et al., 2022) and 86% between 90 and
97 wk of age (Guo et al., 2021). While the 89% laying
rate was reported at an age close to the end of a normal
lay cycle, the 86% laying rate reported beyond 90 wk of
age provided some evidence—albeit based on a very
small sample size—of the possibility for sustaining high
levels of productivity in modern layers to the far end of
the extended lay cycle.
The lowest reported laying rate was 45% at 99 wk of
age (Calik, 2017). Many of the extremely low laying
rates (45−55%) reported had extenuating circumstances
such as high mortality rates, flocks with low productiv-
ity in normal production periods (Calik, 2017), and heri-
tage flocks that were used for comparison to modern
commercial layers (Hanlon et al., 2021). Overall, about
25% of the publications reviewed had laying rates in
extended cycles falling below 70%—the laying rate that
is considered a minimum threshold for commercial via-
bility. The laying rates below 70% were seen mostly at
90 wk of age or beyond in the studies reviewed (Saibaba
et al., 2021).
When only laying rates reported at a specific age are
considered, greater variability was seen at 90 wk and
beyond around the 70% threshold (Figure 2A). Adding
results from the studies that reported laying rate for lon-
ger monitoring periods (by assuming the midpoint as
the reporting age) showed a similar trend (Figure 2B).
Modern commercial layers seem capable of sustaining
laying rates over 70% consistently up to 80 to 82 wk of
age. The variability increased between 82 and 95 wk of
age with productivity levels mostly between 60 and
80%. Though the sample size for 100 wk and beyond was
small, the laying rates were rarely above 70% this late
into a lay cycle.
Reported egg mass ranged between 60 and 70 g—though
trends similar to egg laying rate were observed (Figure 3).
Egg weights were between 60 and 65 g up to 80 wk of age.
Between 81 and 95 wk of age, egg weights were between
62.5 and 67.5 g and increased variability was observed.
Only Anwar and Rahman (2016b)—based on a post-molt
flock—reported egg mass below 60 g (as low as 52 g) at
90 wk of age but this appeared to be a clear outlier. The
mean and median for average egg mass beyond 70 wk of
age was 65 g. Sepehr et al. (2021) reported the highest aver-
age egg weight of 70.11 g at 80 wk of age.
Feed conversion ratio (kilogram of feed required to
produce 1 kg of eggs) was not used as widely as egg lay-
ing rate and egg weight to measure productivity. Over-
all, FCR for lay cycles beyond 70 wk of age were
between 2 and 2.75—with quite a few instances of FCR
above 3 being reported. Trends based on age were not as
clearly seen for FCR as compared to laying rate and egg
weight (Figure 4). Between 70 and 80 wk of age, FCR
was predominantly below 2.5. Feed conversion ratio
below 2—which is close to the feed use efficiency of hens
toward the end of normal Canadian lay cycles—was
reported in only a couple of studies at 80 and 93 wk.
Table S5 in the supplementary file provides a summary
of all of the data that were collected for laying rate, egg
weight, and FCR in this review.
 EXTENDED LAY CYCLES LITERATURE REVIEW
Figure 2. (A) Laying rates reported at a specific age in the literature reviewed. (B) All laying rates reported in the literature reviewed.
The single cycle North Carolina layer performance
tests (Anderson, 2012a, 2015, 2018) did not provide
data separately for the extended lay period. Instead, the
results provided in these test reports (Table 2) provide a
good indication of the overall performance levels that
can be achieved cumulatively across a single-extended
lay cycle (17 wk of age to 85/89 wk of age). Egg laying
rates over 80% and FCR below 2.2 were achieved by a
range of layer breeds in all types of housing over a 68- to
72-wk-long production cycle.
Egg Quality In contrast to productivity, a greater vari-
ety of indicators were used to measure egg quality—
which meant sample sizes were too small for analyzing
many indicators such as egg shape index and dynamic
stiffness. This section discusses the performance
observed for some of the commonly reported egg quality
traits in extended lay cycles.
Eggshell thickness was one of the most commonly
used measures of egg quality—although its effectiveness
as a good indicator of egg quality is questionable (Ketta

and Tumov
a, 2018). Often, uniformity of eggshells is a
better indicator of egg quality than is eggshell thickness,
but the correlation between eggshell quality, breaking
strength, and cracked eggs is well established in
9

literature (Ketta and Tumov
a, 2018). Most of the stud-
ies reported eggshell thicknesses above 0.3 mm—consid-
ered a reasonable standard of good egg quality (Sun et
al., 2012)—but increased variability across studies can
be observed at 80 wk and beyond (Figure 5A). Compar-
ing data between the beginning and end of experiments
did show that eggshell thickness generally decreases fur-
ther into an extended production cycle (Figure 5B).
Over half of the studies reviewed (56%) indicated a
decrease in eggshell thickness with age, whereas 38%
reported increased eggshell thickness. Most of the
instances of increased eggshell thickness were attribut-
able to one particular study (Guo et al., 2021) which
shows the potential benefits of vitamin A and vitamin
K3 supplements on egg quality (observed at 93 and
97 wk of age after being fed supplemented diets at 90 wk
of age).
Variability in eggshell breaking strength clearly
increased at and beyond 80 wk of age (Figure 6A). Egg-
shell breaking strengths were predominantly between
30 N and 40 N at the beginning of extended lay cycles—
this matches the average breaking strength of eggs from
52-wk-old hens (Sun et al., 2012). Breaking strength at
similar levels were reported even at 90 to 100 wk in
10 ARULNATHAN ET AL.

Figure 3. Average egg mass reported in the literature reviewed.

different studies but the increased variability meant that
very low eggshell strength numbers (5−20 N) were also
present. Analyzing studies that provided data on break-
ing strengths both at the beginning and end of extended
lay cycles showed that the decrease in eggshell strength
during an extended lay cycle can be quite significant
(Figure 6B).
Data for cracked eggs (%) had a much smaller sample
size—all 4 studies that had data for this indicator at
both the beginning and the end of the lay cycle indicated
that the proportion of cracked eggs increased signifi-
cantly even within a few weeks (Figure 7A). The 4 North
Carolina layer performance tests included in this review
reported share of cracked eggs over the entire production
cycle. Though the 2012 tests (Anderson, 2012a,b)
showed more variability and higher percentage of
cracked eggs, the most recent studies (Anderson, 2015,
2018) mostly had less than 1% of cracked eggs over the
entire production cycle across different housing systems
and hen strains (Figure 7B).
The final egg quality indicator found commonly in the
literature reviewed was the Haugh unit (HU). The mea-
sure is weighted for a standard large egg weighing 2 oun-
ces (56.7 g). The higher the HU, the better the protein
content and freshness of the egg (Jones, 2012). The US
Department of Agriculture egg grades designates eggs
with HU scores above 72 as AA quality (firm). Eggs
with a HU between 60 and 72 are A grade eggs (reason-
ably firm). Haugh unit scores below 60 indicate poor
internal egg quality (a watery albumen that is thin and
lacking in viscosity) (Jones, 2012).
The data from the literature reviewed showed that,
while HU scores decreased with age, only 3% of the
reported HU scores in extended lay cycles were below 60
(Figures 8A and 8B). Thus, extending lay cycles beyond
70 wk is unlikely to result in eggs with poor internal
(protein content) quality. An assessment of the changes
in quality of eggs laid in commercial farms by Moln
ar et
al. (2016) reflected many of the egg quality trends dis-
cussed herein. The study showed that HU scores

Figure 4. Feed conversion ratios reported for extended lay cycles.

 EXTENDED LAY CYCLES LITERATURE REVIEW 11
Table 2. Cumulative productivity over the entire lay cycle based on the North Carolina layer performance and management tests
reviewed.

White strains— Brown strains— White strains— Brown strains— White strains— Brown strains— Brown strains—
Reference/indicator/scenario conventional conventional enriched enriched cage free cage free range

R24 Age (wk) 17−85

Egg production % 84.6 81.8 83.6 87.6
Egg weight (g) 62 62 61.3 60.7
Feed conversion ratio (FCR) 2.04 2.13 2.13 2.08
R25 Age (wk) 17−89
Egg production % 82.9 90.5 84 78.3 82 81.7 79.1
Egg weight (g) 61.1 60.4 61.5 60.8 62.4 62.3 63.57
Feed conversion ratio (FCR) 2.06 2.18 2.04 2.22 2.27 2.33 2.27
R26 Age (wk) 17−89
Egg production % 86.9 82.9 84 84.5 84 79.6 81.8
Egg weight (g) 60.4 60.7 59.9 61.1 60.3 61.1 63.5
Feed conversion ratio (FCR) 1.92 2.08 2.08 2.13 2 2.17 2.12

decrease at a rate of 0.38/wk after 60 wk of age. Relative
shell weight was found to decrease by 0.02% between 60
and 82 wk, whereas shell thickness became 0.23 mm thin-
ner per week (Moln
ar et al., 2016).
Overall, this literature review is consistent with the
conclusion of Moln
ar et al. (2016) that, while most egg
quality traits decrease with age, the reduction in egg
quality is not significant enough that extending lay
cycles beyond 70 wk of age is not possible. This is further
supported by the cluster analysis of Sirri et al. (2018),
which grouped eggs between 44 and 58 wk of age in one
subgroup and those between 58 and 81 wk of age in
another subgroup based on egg quality traits. All the
egg quality data compiled from the literature reviewed—
including indicators not discussed here—is provided in
the supplementary file (Table S6).
Animal Welfare Consideration of animal welfare out-
comes was much less frequent compared to productivity
and egg quality in the literature reviewed. The only 2
animal welfare indicators for which data were reported
in 3 or more studies were cumulative mortality (12/34
publications reviewed) and tibia breaking strength

Figure 5. (A) Reported eggshell thickness during extended lay cycles in the literature reviewed. (B) Comparing reported eggshell thickness
between the beginning and the end of the experimental period.
12 ARULNATHAN ET AL.
Figure 6. (A) Reported eggshell breaking strength during extended lay cycles. (B) Comparing eggshell breaking strength between beginning
and end of experimental periods.
(5/34 publications reviewed). Over 60% of the reported
cumulative mortality rates were 10% or higher after
80 wk of age (Figure 9A). The median mortality rate for
extended lay cycles in the literature reviewed was
11.7%—compared to an average Canadian mortality
rate of <3% in cage and between 3 and 8% in noncage
housing systems. Despite this, it is important to note
that variability was also significant—as seen in
Figure 9B, which plots the cumulative mortality rates
for various hen strains and housing system at 89 wk of
age reported in (Anderson, 2015, 2018).
The sample sizes were too small to identify possible
age-related trends in tibia breaking strength. Other ani-
mal welfare considerations in the literature (but with not
enough data to inform any analysis) were tibia weight,
tibia length, tibia diameter, feather cover, keel bone devi-
ations, comb wounds, and antioxidative enzymes. A table
summarizing all of the animal welfare data collected is
provided in the supplementary file (Table S7).
Despite the lack of a common subset of indicators to
enable detailed analyses on animal welfare outcomes,
there are several animal welfare concerns that deserve
attention within the context of this review. Increased
incidence of osteoporosis is one of the most referenced
welfare concerns with respect to longer lay cycles (Bain
et al., 2016). This may subsequently lead to bone fragil-
ity and increased susceptibility to bone fracture (Sandi-
lands, 2011). Hens with low egg production or prolonged
intervals between lay at an older age tend to have stron-
ger medullary bone—suggesting a possible (negative)
correlation between egg formation and skeletal integrity
(Dunn et al., 2021). Studying the correlation between
egg production, egg quality, and bone quality, Alfonso-
Carrillo et al. (2021) showed that, while hens with low
productivity and egg quality had better skeletal quality,
there was no clear evidence that productivity/egg qual-
ity and skeletal integrity are necessarily at odds in
extended lay cycles. Both domains can improve if
 EXTENDED LAY CYCLES LITERATURE REVIEW 13

Figure 7. (A) Share of cracked eggs (%) reported at the beginning and end of experimental periods. (B) Cracked eggs data from the four North
Carolina layer performance and management tests—each dot represents the cracked eggs rate reported for a specific strain housed in either caged or
alternate housing systems.

medullary bone with enough calcium reserves is built
early in the hen’s life though delayed onset of lay or high
calcium diets at the pullet phase (Dekalb, 2017).
Increased lay cycle length may also increase the inci-
dence of metabolic diseases, such as fatty liver hemor-
rhagic syndrome (Bain et al., 2016). In addition, lay
cycle length may impact the affective state and behav-
ioral components of animal welfare. Increased disease
incidence, for example, will result in negative contribu-
tions to hen affective state in addition to challenges to
biological health. In contrast, it has been suggested that
fear responses in hens may decrease with age (Hocking
et al., 2001), representing a potential reduced contribu-
tion to negative affective state associated with longer
laying cycles. Behavioral changes may also be expected
as hens continue to age (Sokolowicz et al., 2020).
Feather pecking could pose a major challenge in extend-
ing lay cycles—especially in noncage housing systems
(Bain et al., 2016).
Performance in Extended Lay Cycles Relative to
Housing System and Hen Type Comparing the per-
formance of hens in extended lay cycles by housing sys-
tem type was limited by the small number of studies
using nonconventional cage housing systems, and the
further reduction in sample sizes when trying to account
for age at which results were recorded for commonly
used indicators. Table 3 shows the average cumulative
performance seen in Anderson (2015) and Anderson
(2018)—the only studies in this review that compared
4 housing system types. Further, both of these studies
had lay cycle lengths of 89 wk, used similar breeds of
hens, and had other similarities in study conditions that
allowed for aggregating the cumulative performance
results. These results are compared against the perfor-
mance levels indicated in layer management guides. The
cumulative data for laying rate, FCR, egg weight, and
mortality at 90 wk were collected from the caged and
noncaged management guide of each breed of hen tested
14 ARULNATHAN ET AL.
Figure 8. (A) Reported Haugh unit scores during extended lay cycles. (B) Comparing Haugh unit scores between beginning and end of experi-
mental periods.
in Anderson (2015) and Anderson (2018) and averaged
for comparison.
Overall, conventional cage housing had the best per-
formance for extended lay cycles. The hen daily laying
rate was at least 3% higher in conventional cage housing
(86.2%) at 90 wk compared to enriched colony, free-run,
and free-range systems. Both FCR (2.1) and egg weights
(60.6−61.2 g) were similar for conventional, enriched,
and free-run housing. Free-range housing had the poor-
est performance overall for laying rate (79.9%), FCR
(2.3), and average egg weight (63 g). Mortality rates in
Anderson (2015) and Anderson (2018) were very high in
comparison to the average performance data from man-
agement guides. But for laying rate, egg weight, and
FCR, performance levels seen in the 2 tests were similar
to the management guide averages. Overall, since the
results are based on just 2 studies, no definitive conclu-
sions can be drawn on the differences between housing
system types for extended lay cycles.
The performance levels reported in the literature
reviewed at different points of the late laying phase were
also compared against the management guide averages.
By combining the publications using conventional and
enriched housing into one category (caged housing), lay-
ing hen performance data at 70, 80, and 90 wk of age in
caged systems were generated. The indicators for which
enough data were available were laying rate, egg weight,
HU score, eggshell breaking strength, and cumulative
mortality (Table 4). Due to small sample sizes, similar
estimates could not be generated for noncaged housing.
Hen daily laying rates for caged housing seen in the liter-
ature matched the management guides at 70 wk of age
but were 2.5% points lower at 90 wk of age. By 90 wk of
age, layers are close to reaching the minimum threshold
of 70% rate of lay required for economic viability. Simi-
larly, average egg weight was lower compared to the
management guide at 70 wk, but higher by 90 wk of age.
Eggshell breaking strengths provided in the manage-
ment guides were much higher, with the drop off in egg-
shell strength observed in the data quite significant
between 70 and 90 wk of age. Similarly, HU scores for
eggs from caged housing were much lower than the man-
agement guides. This indicates that the high productiv-
ity levels described in the management guides are much
more achievable than are the predicted egg quality lev-
els. As Sambeek (2011a) mentioned, intensive genetic
selection for extending lay cycles has focused primarily
on maintaining lay persistency and flattening the egg
weight curve. It is likely that focus will increasingly turn
to improving egg quality alongside the higher lay persis-
tency that is now achievable in older laying hens.
Similar to the comparison of housing systems, com-
paring the performance of different breeds of hens was
limited by data availability. Once again, Anderson
 EXTENDED LAY CYCLES LITERATURE REVIEW 15

Figure 9. (A) All cumulative mortality rates reported in the literature reviewed. (B) Cumulative mortality rates for different hen strains and
housing systems at 89 wk of age reported in Anderson (2015) and Anderson (2018)—each dot represents the mortality reported for a specific strain
housed in either caged or alternate housing systems.

(2015) and Anderson (2018) provided some data to com-
pare various white- and brown-type laying hens against
their respective management guides. Table 5 lists the
average laying rate, FCR, egg weight, and cumulative
mortality (from different housing systems) over 90 wk
for the most analyzed white and brown hens in this
review. Performance levels for hen types in these studies
matched the management guides for all indicators
except cumulative mortality. Overall, white-type layers
performed better than brown-type layers in extended
lay cycles in both the literature reviewed and in the man-
agement guides.

Table 3. Comparison of layer performance at 90 wk of age reported in Anderson (2015) and Anderson (2018) to layer management
guide averages.

Laying Feed conversion ratio Egg Mortality

Cumulative (17−90 wk) rate (%) (kg feed/kg of eggs) weight (g) (%)
Conventional 86.2 2.1 60.6 14.7
Enriched 82.8 2.1 60.9 17.3
Management guides (caged) 82.3 2.1 62.6 6.1
Free-run/Aviary 82.4 2.1 61.2 12.9
Free range 79.9 2.3 63.0 4.2
Management guides (alternate housing) 82.9 2.2 62.1 7.6
16 ARULNATHAN ET AL.

Table 4. Performance of laying hens at 70, 80, and 90 wk of age in caged systems in the literature reviewed compared against average
management guide data.

Management Management guides

Indicator Age (wk) Cage guides (caged) (alternate housing)
Laying rate (%) 70 83.2 83.8 86.4
80 77.5 78.7 80.8
90 71.3 73.7 76.4
Egg weight (g) 70 63.4 64.4 64.6
80 65.1 64.8 64.9
90 65.8 65.5 65.1
HU score 70 76.6 82.7 81.0
80 78.0 81.7 80.4
90 71.0 81.3 80.4
Eggshell breaking strength (N) 70 38.0 40.4 40.8
80 34.8 39.7 40.5
90 31.0 39.4 40.3
*Cumulative mortality (%) 70 2.8 4.0 4.7
80 3.5 4.9 6.1
90 4.7 6.1 7.6
*
Mortality only for experiment period.

Figure 10. (A) Comparing the feed costs and egg income of different housing systems. (B) Comparing feed costs and egg income of conventional
housing in lay cycles ending at 85, 89, and 109 wk.
 EXTENDED LAY CYCLES LITERATURE REVIEW 17

Management
Economic Performance Profitability is crucial in

guides

6.86
6.32
determining the likelihood of Canadian farmers adopt-

6.0
7.1
6.3
7.5
7.8
6.5
4.6
6.8
7.3
6.3
6.7
Cumulative mortality (%)
ing extended lay cycles. Very little economic information
related to extended lay cycles was obtained in this litera-
ture review. Except for the North Carolina layer perfor-
and Anderson (2018)

mance tests (Anderson, 2012a,b, 2015, 2018), none of

Anderson (2015)

the publications considered provided any economic

data. This is a major gap in literature that requires

16.67
13.47
26.7
16.9
17.2
14.6
18.1

10.1
15.8
6.6

21.8

10.3
9.4
addressing. However, the feed cost and egg income data
from the North Carolina tests provided an indication
that profitability is achievable with extended lay cycles
in all housing systems (Figures 10A and 10B). Compar-
ing egg income and feed costs for conventional housing
Management

at 85, 89, and 109 wk showed that extending lay cycles

guides
61.70
62.00
62.20
61.15
61.60

63.50
63.00

63.40
61.73
63.30
clearly has an inflexion point (between 89 and 109 wk)
−
−

beyond which the increased costs of keeping hens in lay

is not compensated by egg income due to reduced lay,
Egg weight (g)

higher mortality, and poor feed conversion efficiencies.

and Anderson (2018)

Further, these studies (Anderson, 2012a,b, 2015, 2018)

Anderson (2015)
Table 5. Comparison of popular hen breeds based on cumulative performance data from Anderson (2015) and Anderson (2018).

did not account for eggs discarded at grading. The

increased loss in egg quality seen around 100 wk of age
60.13
59.32
60.35
62.64
61.96
60.02
58.31
61.74
61.58
61.19
62.09
60.74
60.98

could potentially further reduce profitability. However,

since the 4 studies from which economic data were
sourced were conducted in different years (which influ-
ences the feed costs and the egg prices used to calculate
egg income), these trends need further exploration
Management

before any definitive conclusions can be supported.

guides
1.99
2.03
2.11
2.01
2.05
2.15
2.10
2.27
2.15
2.14
2.26
2.06
2.18

Meat Characteristics Eggs are not the only valuable

FCR (kg feed/kg of eggs)

output of commercial egg production. The spent hens

represent a valuable co-product that can be used subse-
quently for human consumption or rendering for animal
and Anderson (2018)

feed. Only one study—Yalcin et al. (2020)—identified in

Anderson (2015)

this review addressed how the quality and characteris-

tics of meat can change as a result of a longer lay cycle.
1.97
2.06
2.06
2.02
2.07
2.13
2.25
2.13
2.10
2.17
2.17
2.05
2.17

Yalcin et al. (2020) found that extension of lay cycles

from 75 to 108 wk caused a reduction in the share of
breast and thigh meat, affected the appearance of the
meat (darker/yellower), and increased fat content in
breast meat. Despite the decrease in quality, Yalcin et
Management

al. (2020) concluded that the meat quality in spent hens

guides
82.80
83.40
83.85
83.20
81.95
84.50
81.20
80.10
80.75
85.10
80.80
83.28
81.59

at 108 wk was still good enough for further use.

Laying rate (%)

CONCLUSIONS
and Anderson (2018)
Anderson (2015)

Extending lay cycles beyond current Canadian norms

of 50 to 52 wk is a potentially important strategy in
85.33
83.14
86.78
85.00
83.68
79.27
85.22
84.20
83.83
82.03
84.73
83.86
84.00

improving some sustainability outcomes—but the pro-

ductivity, egg quality, and animal welfare outcomes
associated with this practice are not sufficiently estab-
lished to enable definitively estimating net impacts and
benefits. In light of there having been no systematic
Layer breed (cumulative—

review of literature on extended lay cycles published to

Brown-type hens—overall
White type hens—overall

date, the objectives of the current analysis were to iden-

Hy-Line Silver Brown

tify relevant literature, establish performance levels

Lohmann LSL Lite

Lohmann LB-Lite
H&N Nick Chick

under different lay cycle lengths, and identify key strate-

Hy-Line Brown
Bovans Brown
Bovans White

Hy-Line W-36
Dekalb White
Shaver White
90 wk of age)

gies and management practices that can support suc-

ISA Brown

cessfully extending lay cycles. A wide range of lay cycle

lengths (end of lay at 72−119 wk) and hen breeds were
represented in the review but most of the analyses were
18 ARULNATHAN ET AL.
restricted to conventional cage housing systems. Several
feed-related interventions such as high-protein diets,
probiotics supplements and vitamin A and K3 supple-
ments identified through this review have the potential
to positively influence productivity and, especially, egg
quality during the late laying phase. Alternate feed
delivery methods such as time-restricted feeding and
split-feeding also have the potential to improve perfor-
mance in aged laying hens. Overall, productivity was
acceptable beyond 70 wk of age with hens close to falling
below 70% laying rate only at 100 wk and beyond. Per-
formance on most egg quality traits and animal welfare
indicators also decline with age. However, like produc-
tivity, performance is still at acceptable levels well past
70 wk of age. There was increased variability beyond
»80 wk of age across most productivity, egg quality and
welfare indicators—which highlights that there might
be factors such as flock management and specific dietary
and rearing practices during the pullet phase that can
help realize better performance during an extended lay
cycle. Lack of sufficient data restricted any detailed
analysis of layer hen performance in extended cycles
according to housing system type and layer type—
although there are (inconclusive) indications that hens
in caged housing and white laying hens are better than
hens in noncaged housing and brown-type layers. Eco-
nomic data were also limited but reinforced the observa-
tions seen in the productivity and egg quality metrics
that there could potentially be a tipping point beyond
90 wk of age after which persisting with an extended lay
cycle might not be economically worthwhile. Based on
the results of this review, we make the following recom-
mendations as to important next steps.
 With the on-going transition to nonconventional
housing in countries like Canada, more studies on
performance of extended lay cycles in alternate hous-
ing systems—especially free run and free range—is
required.
 Identification of a common subset of egg quality and
animal welfare indicators is required with respect to
extended lay cycles to enable better comparison and
aggregation of data for analysis.
 More research on potential animal welfare impacts
such as pecking wounds, feather loss, and stress
events, as well as economic outcomes associated with
extended lay cycles is required.
 Given the increased variability observed with age and
the possibility that extending lay cycles too long
might negatively influence sustainability outcomes, it
is important to generate more publicly available
information on the performance of laying hens at 90
wk and beyond.
 Pullet-specific management practices and strategies
were not covered in this review due to possible fail-
ures in designing the search strategy. Pullet rearing is
one of the most important aspects of extending lay
cycles and, hence, a systematic literature review simi-
lar current analysis focused exclusively on the pullet-
rearing phase would be useful.
 Further research on optimizing strategies with the
potential to positively influence layer performance,
such as time-restricted feeding, increasing bioavail-
ability of calcium, and use of high-protein and probi-
otic-supplemented diets, is required. Other feed-
related aspects such as changes in energy and amino
acid provisions during the late laying phase also
require analysis.
ACKNOWLEDGMENTS
This work was supported with funding through
research grants from both Egg Farmers of Canada
(EFC) and Egg Farmers of Alberta (EFA)/Results
Driven Agricultural Research (RDAR).
DISCLOSURES
The authors declare that they have no known compet-
ing financial interests or personal relationships that
could have appeared to influence the work reported in
this paper.
SUPPLEMENTARY MATERIALS
Supplementary material associated with this article
can be found in the online version at doi:10.1016/j.
psj.2024.103475.
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