Attention deficits linked with proclivity to
royalsocietypublishing.org/journal/rspb
Research
Cite this article: Barack DL, Ludwig VU,
Parodi F, Ahmed N, Brannon EM,
Ramakrishnan A, Platt ML. 2024 Attention
deficits linked with proclivity to explore while
foraging. Proc. R. Soc. B 291: 20222584.
https://doi.org/10.1098/rspb.2022.2584
Downloaded from https://royalsocietypublishing.org/ on 01 March 2024
Received: 7 April 2023
Accepted: 12 January 2024
Subject Category:
Neuroscience and cognition
Subject Areas:
ecology, health and disease and epidemiology,
cognition
Keywords:
attention deficit hyperactivity disorder, ADHD,
foraging, behaviour, adaptation,
behavioural biomarker
Authors for correspondence:
David L. Barack
e-mail: dbarack@gmail.com
Arjun Ramakrishnan
e-mail: arjun.ramakrishnan@gmail.com
† but potentially depleting option or disengage and search for something that
Co-first-authors.
Electronic supplementary material is available
online at https://doi.org/10.6084/m9.figshare.
c.7049958.
Electronic supplementary material is available
online at https://doi.org/10.6084/m9.figshare.
c.7049958.
explore while foraging
David L. Barack1,2,†, Vera U. Ludwig1,5,†, Felipe Parodi1, Nuwar Ahmed3,
Elizabeth M. Brannon3, Arjun Ramakrishnan6 and Michael L. Platt1,3,4,5
1
Department of Neuroscience, Perelman School of Medicine, 2Department of Philosophy, 3Department of
Psychology, 4Department of Marketing, Wharton School, and 5University of Pennsylvania, PA 19104, USA
6
Department of Biological Sciences and Bioengineering and Mehta Family Centre for Engineering in Medicine,
Indian Institute of Technology, Kanpur 208016, India
DLB, 0000-0001-7131-4638; MLP, 0000-0003-3912-8821
All mobile organisms forage for resources, choosing how and when to
search for new opportunities by comparing current returns with the average
for the environment. In humans, nomadic lifestyles favouring exploration
have been associated with genetic mutations implicated in attention deficit
hyperactivity disorder (ADHD), inviting the hypothesis that this condition
may impact foraging decisions in the general population. Here we tested
this pre-registered hypothesis by examining how human participants
collected resources in an online foraging task. On every trial, participants
chose either to continue to collect rewards from a depleting patch
of resources or to replenish the patch. Participants also completed a
well-validated ADHD self-report screening assessment at the end of
sessions. Participants departed resource patches sooner when travel times
between patches were shorter than when they were longer, as predicted
by optimal foraging theory. Participants whose scores on the ADHD
scale crossed the threshold for a positive screen departed patches signifi-
cantly sooner than participants who did not meet this criterion.
Participants meeting this threshold for ADHD also achieved higher
reward rates than individuals who did not. Our findings suggest that
ADHD attributes may confer foraging advantages in some environments
and invite the possibility that this condition may reflect an adaptation
favouring exploration over exploitation.
1. Introduction
All mobile organisms forage for resources such as food, water and mates [1,2].
Foraging for food that is difficult to find or extract has been hypothesized to be
a major driver of the evolution of intelligence [3–5]. Foraging models have
recently been applied to human behaviour in multiple disciplines including
cognitive psychology [6], cognitive neuroscience [7,8] and computer science
[9,10], validating an approach grounded in mathematical ecology to study
the computations and mechanisms that evolved to regulate the tradeoff
between exploitation and exploration [11].
Patch foraging is the iterated accept-or-reject decision to stick with a known
might be better [12]. This description characterizes a wide array of decision
contexts, including when to search for resources like food, water, minerals
or sexual encounters; internal searches through concepts [13], memory [14] or
strategies [15,16]; or searches for abstract resources such as for information
[9,17,18] or reputation. These decisions can be solved using simple algorithms
derived from optimal foraging theory [1], such as the Marginal Value Theorem
(MVT) [19]. These algorithms dictate that individuals should leave a depleting
resource patch when local intake rates fall below the average for the environ-
ment. Hundreds of species tested—from bees [20] to birds [21] to monkeys
[22] to humans [23]—either quantitatively or qualitatively behave in accordance
© 2024 The Author(s) Published by the Royal Society. All rights reserved.
 with the predictions of such models, suggesting that evol-
ution long ago settled on a near-optimal solution to the
challenge of determining when to abandon a depleting
resource and search for a new one.
Whether searching for external, internal or abstract
resources, foraging requires balancing a tradeoff between
exploring for new options and exploiting known ones.
Attention deficit hyperactivity disorder (ADHD) is a neurode-
velopmental disorder affecting 11% of children [24] and 4.25%
of adults [25]. Symptoms include distractibility as well as rest-
lessness and excessive movement [26]. These characteristics
may impact foraging behaviour by promoting a tendency to
abandon the current resource patch to search for a new one
[11]. This tendency to explore while foraging might extend to
other behaviours such as cycling more frequently between
information sources in the classroom or sources of stimulation
in the home environment [11]. Human populations, such as
the Ariaal tribe in Africa, that live nomadic lifestyles favouring
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exploration are characterized by genetic mutations implicated
in ADHD [27], consistent with the hypothesis that this con-
dition may impact foraging decisions in other populations.
In addition, the neurological and genetic basis of ADHD
has a well-established noradrenergic component [28–30]
linked to task focus [31] and foraging behaviour [32]. Given
the link between distractible, impulsive behaviour and atypical
activity in noradrenergic circuitry, we predict that indivi-
duals with ADHD or ADHD-like phenotypes will both leave
patches earlier than predicted by foraging models like
the MVT and also earlier than individuals lacking these
phenotypical characteristics.
We sought to test these predictions empirically by study-
ing the relationship between foraging behaviour and
ADHD self-report scores in a large population of adults. We
measured ADHD-like symptoms using the Optimal RiskSLIM
DSM-5 ASRS Screening Scale (ASRS) [33]. We designed
an online incentive-compatible virtual berry-picking game
to assess the impact of ADHD symptoms on foraging
behaviour. We tested the pre-registered prediction that stron-
ger self-reported ADHD symptoms would be associated
with shorter patch residence times, less optimal behaviour
and lower cumulative rewards in our online foraging task.
In our sample of 457 participants, time in patch, travel
time and harvested reward all influenced the decision to
continue to forage at a patch or to depart and travel to a
new patch. Overall, participants stayed longer in patches
than predicted by MVT. We found that participants with
elevated scores on the self-report ADHD scale exhibited
shorter patch residence times than those whose scores were
not elevated. Surprisingly, ADHD screen-positive participants
also achieved higher reward rates and their patch-leaving
decisions were more closely aligned with those predicted by
the MVT, although they still stayed longer in patches than
the MVT predicts.
ADHD is thus associated with shorter patch dwell times
and higher reward rates in the online foraging environment
implemented here. These findings are consistent with systema-
tic amplification of activity within neural circuits involved in
exploration, such as the default mode network (DMN)
[12,34], and changes in levels of the neuromodulator norepi-
nephrine broadcast from the locus coeruleus [31,35,36], both
implicated in individuals with ADHD [28–30,37]. Our findings
resonate with field studies highlighting a potential adaptive fit
between population variation in genes linked to ADHD and
nomadic lifestyles that favour exploration [27], endorsing 2
approaches to variation in human behaviour grounded in
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evolutionary medicine [38] and computational psychiatry [39].
2. Methods and materials
(a) Task
We recruited a representative US sample (n = 506) via Prolific
(see https://www.prolific.com/) to complete a virtual
patchy foraging task programmed in Javascript. All methods
were approved by the Internal Review Board of the University
of Pennsylvania. After preprocessing, 49 individuals were
excluded from final analyses for the following criteria. First,
Proc. R. Soc. B 291: 20222584
they either failed pre-defined survey attention checks or did
not complete every survey. Further, participants were excluded
if they completed less than 25 trials (i.e. an exploit or explore
decision) throughout the session, not counting incomplete
patches (see below). Finally, we excluded participants that
were believed to have misunderstood the instructions given
their comments (e.g. ‘I don’t know if I understood it’ or ‘I did
not understand until the last round’). Note that exclusion cri-
teria were slightly changed compared to the pre-registered
criteria (see below). Of the final 457 participants (µage =
45.63 y.o., σage = 16.09), 232 identified as ‘male’, 217 identified
as ‘female’, 5 identified as ‘other’, and 3 chose not to provide
their gender. The final sample was composed of 71% (n =
326) White Caucasian (non-Hispanic), 14% (64) African-Amer-
ican or Black, 8% (38) Asian, 4% (19) Spanish, Hispanic or
LatinX, <1% (3) American Indian or Alaskan Native, and less
than 2% (7) unreported.
In the task, a ‘valid patch’ begins with an exploit decision,
in which the user collects berries, and ends with an explore
decision, in which the user travels to the next patch. Note
that any patches where the participant immediately explored
was considered a valid decision, too, and thus included in the
analysis. Any ‘incomplete patch’ that did not end in the user
leaving the patch (explore) was removed. These incomplete
patches occurred at the end of session blocks in which the
current block timed out after 2 min.
Participants were sent to the training and task via a link in
Qualtrics (see https://www.qualtrics.com/uk/?rid=ip&prev-
site=en&newsite=uk&geo=BE&geomatch=uk) after completing
an informed consent form and several questionnaires. Before
starting, participants received detailed written instructions on
the task and watched a video demonstration. To familiarize
themselves with the task, they also completed a 1 min training
(30 sec per block type) before starting the actual experiment.
The instructions prior to the training were (emphasis in italics
were coloured in original):
‘In this experiment, your goal is to collect as many berries as you
can in the amount of time you are given.
First, hover over the centre box until it fills up completely. You
may have to wiggle your cursor once for the centre box to
recognize your mouse.
Then, on each trial, you will either choose to stay at the
bush you are at or leave the current bush and travel to a
fresh bush.
To pick berries from the current bush, hover your cursor over
the bush next to the centre box.
The number of berries you pick at each attempt will be
indicated next to the bush.
 As you keep picking from the same bush repeatedly, the
number of berries you pick at each attempt will go down
as you will be depleting this bush.
To travel to the next bush, mouse over the box on the opposite
side of the bush.
You can choose to leave the current bush to travel to a fresh
bush at any time. When you do that, you will need to wait
while you travel to the next bush.
However, once you arrive at the new bush, the amount of ber-
ries you pick at each harvest will reset to the initial value.
You will play different blocks in this game. The environment
will change between these blocks.
There is an unlimited number of bushes in the environment,
but a finite amount of time. Try to collect as many berries
as you can!’
After completing the experiment, participants returned to
Qualtrics to finish further questionnaires. Foraging data were
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stored on a secure AWS server.
During the task, participants collected as many berries as
possible in 8 min (4 blocks, 2 min each). The reward function
for the task started at 7 plus Gaussian noise, ε, for both long
and short travel time patches for all patches:
r0 ¼ 7 þ 1,
where ε ∼ N(0, 0.25) and then, for each choice to harvest
reward, ri, decremented by 0.5 plus noise:
ri ¼ ri1  0:5 þ 1,
where again ε ∼ N(0, 0.25). At the end of the experiment, partici-
pants received 0.3226 cent per berry collected (up to $3 in total)
in addition to their base participation compensation of $4.
Participants were randomly assigned to one of two orders
of travel times by block: order 1 [block 1: 1 s travel time, block
2: 5 s, block 3: 1 s, block 4: 5 s] or order 2 [block 1: 5 s travel
time, block 2: 1 s, block 3: 5 s, block 4: 1 s]. Each block lasted
for 2 min. Participants foraged through as many patches as
they wished and were able to in those 2 min; patches at the
end of each block that did not terminate with a leave decision
were removed from analysis. Participants completed an
average of 119.95 ± 1.79 trials on average during the task.
The maximum number of trials was 179 and the minimum
was 25 trials, for a range of 154 trials.
(b) Survey
Participants completed the Optimal RiskSLIM DSM-5 ASRS
Screening Scale [33], consisting of 6 items. The items were:
How often do you have difficulty concentrating on what people
are saying to you even when they are speaking to you directly?
How often do you leave your seat in meetings or other situations
in which you are expected to remain seated?
How often do you have difficulty unwinding and relaxing when
you have time to yourself?
When you’re in a conversation, how often do you find yourself
finishing the sentences of the people you are talking to before
they can finish them themselves?
How often do you put things off until the last minute?
How often do you depend on others to keep your life in order
and attend to details?
Response options were never, rarely, sometimes, often, and
very often. To score the survey, a complex, linear, proprietary
scoring function was received from its authors [33]. We are
not at liberty to share the method here, but researchers can 3
obtain it after completing a use agreement with the scale
royalsocietypublishing.org/journal/rspb
authors (see https://license.tov.med.nyu.edu/product/asrs–
dsm-5 for instructions). This is currently free for all academic
purposes. If there are issues obtaining the scoring method,
researchers could revert to scoring the scale using a simple
sum (see electronic supplementary material).
The ADHD self-report symptom checklist from Ustun et al.
[33] used a machine learning algorithm, the risk-calibrated
super-sparse linear integer model (RiskSLIM), to generate an
optimal scoring algorithm for the ADHD Self-Report Screening
Scale survey based on the updated DSM-5 criteria for ADHD.
The scale was validated by comparison to a managed care
sample of ADHD diagnosed adults (n = 218), a household
Proc. R. Soc. B 291: 20222584
sample from the National Comorbidity Survey Replication
consisting of a face-to-face survey and follow-up (n = 119)
and an NYU-Langone sample (n = 300). At the recommended
scoring threshold (≥14) for a positive screen, the ASRS has
good sensitivity (91.4%), very good specificity (96.0%), an
area under curve (AUC) of 0.94, with 67.3% of screen-positive
participants meeting DSM-V criteria for ADHD among a com-
bined cohort from the National Comorbidity Survey-
Replication and a managed care sample, along with good sen-
sitivity (91.9%), moderate specificity (74.0%), an AUC of 0.83
and a proportion of 82.8% of screen-positive participants
who met DSM-V criteria for ADHD among a clinical sample
from NYU-Langone Medical Center [33]. The scale has been
further validated by other groups [40–44].
(c) Preregistration
Our design, hypotheses and analyses were preregistered: see
https://aspredicted.org/blind.php?x=KLK_XLL.
We made a few justified changes and additions to the pre-
registered analysis approach. We analysed total reward per
patch and reward rates in both conditions (short versus long
travel time) rather than total reward over the entire experiment,
because foraging theory predicts differences between patch
types. We calculated optimal behaviour and deviations from
optimal behaviour in a way that differed from the preregistered
version because we considered the approach by Cowie [45] to
be more accurate than the pre-registered proposal. Specifically,
Cowie’s method presents a first principles way to calculate
optimal behaviour. In the binomial regression analysis includ-
ing ASRS, we included ‘cumulative reward in patch’ as
an additional predictor because we considered it relevant to
the model. We did not include trait and state anxiety or positive
and negative emotions as control variables. The categorization
of participants into ADHD-positive and ADHD-negative
was not preregistered. However, since the ASRS scale uses a
complex linear scoring algorithm to classify participants as
positive or negative, it made sense to use this qualitative and
clinically meaningful distinction between participants in our
analysis. ‘Give up time’ ( pre-registration) is referred to as
‘time in patch’ or ‘patch residence time’ in the current manu-
script, as we considered these to be more accurate terms. The
analysis of the link between ASRS with creativity and mind
wandering will be reported separately for brevity here. Finally,
exclusion criteria for participants were slightly improved.
For example, in addition to the pre-registered criteria, we
also excluded participants who commented that they did not
understand the task. Moreover, trials belonging to incomplete
patches at the end of each session were excluded before
applying the ≥25 trials criterion.
 (d) Data analysis
All data were analysed in MATLAB (Mathworks, Natick,
MA) using custom software. For tests involving comparisons
across 1 s and 5 s travel times, 1 subject was rejected who
never completed a 5 s travel time patch, leaving n = 456.
For patch residence time comparisons, we averaged the
median patch residence time across subjects, to control for out-
liers. We used within-subject paired t-tests to compare the
average patch residence time for 1 s and 5 s travel time patches.
Time in patch was the amount of time from the start of the first
trial in a patch to the decision to depart a patch.
For reward-related comparisons, we used both cumulative
reward (which previous work has shown to be the computa-
tionally relevant variable for decisions to harvest rewards in
patch [46]) and reward rate (the classic currency for foraging
choices in optimal foraging theory [1]). Cumulative reward
was defined as the running sum of berries collected within a
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patch, and reward rate was defined by dividing cumulative
reward by the cumulative time in patch. We used within-subject
paired t-tests between travel times to compare cumulative
rewards and reward rates. We also examined differences in
cumulative reward across all blocks in the entire session for
ADHD screen-positive and screen-negative participants.
To assess the effects of patch number in session, we first
segregated total time in patch and reward rate on patch
leave trials by travel time block (1 s or 5 s). We then separately
regressed time in patch and reward rate upon departing
patches against patch number in session using ordinary least
squares (OLS) regressions. Patch number in session was
defined as the patch number in the concatenated blocks
segregated by travel time.
ADHD self-report scale (ASRS) scores were correlated
with both time in patch and reward rates. To determine the
influence of ASRS on time in patch, we used a mixed-effects
linear regression, with random effects of the side of the screen
of the choice target, patch number in session, and participant
identity and fixed effects of travel time, cumulative reward
and ASRS score, with all interactions, and fixed effects
of nuisance covariates age, income and gender, with all
interactions. In Wilkinson notation,
Time in patch
CumulativeReward  TravelTime
 ASRSScore þ Age  Gender  Income
þ (Side þ PatchNumberinSessionjParticipantNumber),
which models patch number in session as a random slope
and participant identity as a random intercept. To determine
the influence of ASRS on decisions to leave patches, we used
a mixed-effects binomial regression, with random effects of
the side of the screen of the choice target, patch number
in session, and participant identity and fixed effects of
time in patch, travel time, cumulative reward and ASRS
score, with all interactions, and fixed effects of nuisance
covariates age, income and gender, with all interactions.
In Wilkinson notation,
~
ProbabilityChooseLeaveTimeInPatch
 CumulativeReward  TravelTime  ASRSScore
þ Age  Gender  Income
þ (Side þ PatchNumberinSessionjParticipantNumber),
which again models patch number in session as a random
slope and participant identity as a random intercept. Finally,
we used Student’s t-tests to compare ADHD screen-positive 4
with screen-negative time in patch, and reward rate
royalsocietypublishing.org/journal/rspb
distributions for the two groups were compared using a
Kolmogorov-Smirnov test.
(e) Optimal behaviour
First, an exponential curve was fit to the cumulative intake
curve defined by the gain function ( fit function in MATLAB).
The gain function g(t) for time in patch t is well-described by
an exponential curve, which can be used to solve from first
principles the optimal leave time [cf. 45]:
g ¼ b1 (1  eb2 t ): ð2:1Þ
Proc. R. Soc. B 291: 20222584
While this continuous and differentiable function is at
odds with the discrete real series of rewards, describing the
series of rewards as an exponential permits a first principles
solution to the optimal patch stay time. To find the optimal
leave times, the first derivative of g(t) is set equal to the
gain function divided by the average travel time and time
in patch:
dg g
¼ b1 b2 eb2 t ¼ , ð2:2Þ
dt tt þ ^t
for travel time tt and optimal leave time ^t. Substituting (2.1) in
for g and rearranging terms yields
^
eb2 t  b2^t ¼ b2 tt þ 1: ð2:3Þ
We next used the McLaurin expansion for e, eliminated
constants, dropped the fourth-order and higher terms,
subtracted out the first-order term and rearranged to arrive at
6tt
3^t2 þ b2 ^t3  ¼ 0: ð2:4Þ
b2
We used the best-fit β2 from our exponential fit (2.1) to the
series of average returns. Then, using the average (tt = 3 sec),
short (tt = 1 sec) and long (tt = 5 sec) travel times in turn, we
determined the optimal travel time ^t by solving (2.4) with
numerical methods ( fmincon in MATLAB subject to nonlinear
constraints determined by the McLaurin expansion).
3. Results
(a) Foraging task and behaviour
Adults (n = 457) performed an online foraging task (figure 1).
Participants were recruited via Prolific and were a representa-
tive US sample (47% female, average age = 46y). Participants
were instructed to collect as many berries as possible in 8 min
(4 blocks, 2 min each). Participants first initiated a trial by
mousing over an intertrial interval (ITI) box. After initiating
a trial, participants chose between mousing over an image
of a bush to stay and collect rewards at a given patch, in
which payoffs declined with each successive choice, or mous-
ing over an empty-sided box to travel to a new patch, which
took time. Each participant was tested in two conditions that
differed only in the time it took to ‘travel’ to the next bush
(either 1 sec or 5 sec). Travel time was indicated by the
visual distance between bushes (see figure 1, inset). The
order of conditions was counterbalanced across participants.
After deciding to depart a patch, bushes moved vertically
down the screen, simulating travel, and the participants
 stay 5
move cursor to
current bush:

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collect berries
at current patch
+6.75
move cursor to ITI
box to start trial upon cursor contact: upon cursor contact: start new trial in the same
ITI box borders turn collected reward is patch by moving cursor to
dark green after 1 s: displayed for 1 s ITI box
ITI box turns solid green leave
stay or leave?
move cursor
to empty side
travel time to next patch is indicated by distance of box: move to
patches to each other throughout each block a fresh patch
1 s travel time 5 s travel time
2 blocks of 2 min each 2 blocks of 2 min each
upon cursor contact with box: start new trial in the next
bushes move down for 1 s patch by moving cursor to
or 5 s, depending on block, ITI box; current patch is now

Proc. R. Soc. B 291: 20222584

simulating travel on the opposite side
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Figure 1. Virtual patch foraging task. Participants began each trial by moving their mouse cursor over an intertrial interval (ITI) box. After 1 s delay, the ITI box turned
green. Participants next decided to stay at a given patch and gather points (’exploit’ decision) or to replenish the patch in exchange for a travel time delay (’explore’
decision). Travel times were either 1 s or 5 s, in blocks. Inset: screenshots of the two foraging environments, with 1 s travel times on the left and 5 s on the right.

(a) 27 (b)
35
26 34

25 33
median total time in patch (s)

rewards collected (arb. units)

32
24
31
23
30
22
29
21
28
20 27
19 26
18 25
1 5 1 5
travel time (s) travel time (s)
Figure 2. Human participants’ (n = 457) behaviour on the virtual patch foraging task. (a) Mean median total time in patch (s) for the 1 s travel time (left) and 5 s
(right). (b) Mean rewards harvested for 1 s travel time (left) and 5 s (right).

faced a new set of decisions between staying in a patch and time environments (RRlong,optimal = 2.2256, p < 1 × 10−119,
departing, with the choice options—the bush and the t (d.f. = 455) = −32.4675). This suboptimal behaviour resulted
empty box switching their sides on the screen. from participants staying for too long in each patch (time
Consistent with predictions from foraging theory’s MVT in patch TiP; short patches: TiPshort = 19.5430 ± 0.1360 s,
[19], participants abandoned patches in a short travel time TiPshort,optimal = 6.0603 s; long patches: TiPlong = 25.5998 ±
(1 sec) environment earlier than they left patches in a long 8039 s, TiPlong,optimal = 12.9049 s), as observed in prior studies
travel time (5 sec) environment (paired t-test on median patch in other animals including nonhuman primates [22,47–50].
residence time by subject, p < 1 × 10−17, t (d.f. = 455) = −9.1942; In addition to these travel time and reward rate effects,
figure 2a). Participants earned significantly more rewards exploratory data analysis uncovered an effect of patch
on average in long travel time environments (paired t-test, p < number in session. Participants performed up to four blocks
1 × 10−44, t (d.f. = 455) = −15.7081; figure 2b), but longer resi- of patches, starting with a block of 1 s patches, then a block
dence times in long travel time patches did not yield of 5 s patches, then 1 s and then 5 s again, or the reverse
significantly higher average reward rates (RR; average order (5, 1, 5, 1; see §2a). Some participants showed an effect
RRshort = 1.4597 ± 0.0271 points s−1, average RRlong = 1.4765 ± of patch number in session on patch residence time and
0.0242 points s−1; paired t-test, p > 0.22, t (d.f. = 455) = −1.2113). reward rate. Segregating by travel time, some subjects
Average reward rates were significantly less than the theor- showed an effect of patch number in session for patch residence
etically optimal reward rates in both short travel time time (1 s patches: OLS, 113/457 participants significant ( p <
environments (RRshort,optimal = 2.6118 points s−1; one-sample 0.05) slope, 74 negative; 5 s patches: OLS, 85/457 participants
t-test, p < 1 × 10−169, t (df = 456) = −45.0179) and long travel significant slope, 53 negative; 168 / 457 unique participants
 (a) participant no. 4 participant no. 270 6
30 26
short travel time patch

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long travel time patch 24
total time in patch (s) 25 22

total time in patch (s)

20
20
18
15 16
14
10 12
10
5
8
6

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0 2 4 6 8 10 12 14 16 18 20 0 2 4 6 8 10 12 14 16
patch number in session patch number in session

(b) participant no. 4 participant no. 270

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3.0 3.4

reward rate on leaving (berries s–1)

reward rate on leaving (berries s–1)

3.2
2.5
3.0
2.0 2.8
2.6
1.5
2.4
1.0 2.2
2.0
0.5 short travel time patch
long travel time patch 1.8
1.6
0 2 4 6 8 10 12 14 16 18 20 0 2 4 6 8 10 12 14 16
patch number in session patch number in session
Figure 3. Participants’ patch residence time and reward rate on leaving a patch as a function of patch number in session, concatenated across blocks for 1 s and 5 s
travel times separately. (a) Total time in patch (s) versus patch number in session, for 1 s (green) and 5 s (red) travel times. Participant no. 4 (left) did not show a
patch number in session effect, whereas participant no. 270 (right) showed a steep decline in total time in patch as a function of patch number in session for both
travel times. (b) Reward rate on patch leave (berries s−1) versus patch number in session for 1 s (green) and 5 s (red) travel times. Participant no. 4 (left) did not
show a change in reward rates as a function of patch number in session, whereas participant no. 270 (right) did, increasing their reward rate.

showed some effect). Two example participants are shown in in session, patch residence time tended to decrease and
figure 3a; the left participant (no. 4) did not show an effect of reward rates increase across sessions; in short, participants
patch number in session (1 s patches: OLS, βslope = 0.1741 ± tended to perform better as they progressed through the task.
0.1490 s/patch number, p > 0.25; 5 s patches: OLS, βslope =
0.1860 ± 0.4653 s/patch number, p > 0.69), whereas the
participant on the right (no. 270) did (1 s patches: OLS, (b) ADHD Self-Report Scale and foraging
βslope = −0.8646 ± 0.1550 s/patch number, p < 0.001; 5 s We next examined how each of the task variables was related
patches: OLS, βslope = −1.5769 ± 0.2684 s/patch number, p < to participants’ ADHD Self-Report Scale scores (ASRS [33]).
0.001). For reward rate, some subjects showed an effect of We first tested for patch-level effects of ASRS by performing
patch number in session (1 s patches: OLS, 131/457 partici- a mixed-effects linear regression of time in patch (n = 10 353
pants significant ( p < 0.05) slope, 126 positive; 5 s patches: patches) against a random effect of patch number in session
OLS, 96/457 participants significant slope, 84 positive; and participant, fixed effects of travel time, cumulative
187/457 unique participants showed some effect). These reward and ASRS scores and all interactions, and a number
same two participants illustrate these effects on reward rate of nuisance covariates including age, gender and income
(figure 3b), with the participant on the left (no. 4) showing and all interactions (see §2d). We reasoned that because of
no effect of patch number in session on reward rate (1 s patches: differences across subjects regarding patch numbers in ses-
OLS, βslope = 0.0114 ± 0.0291 (berries s−1)/patch number, p > sion, covariate should be treated as a random effect. All
0.7; 5 s patches: OLS, βslope = 0.0129 ± 0.0163 (berries s−1)/ fixed effect covariates were z-scored. We uncovered a main
patch number, p > 0.45), unlike the participant on the right effect of cumulative reward ( p ∼ 0, Bonferroni-corrected),
(no. 270) who did (1 s patches: OLS, βslope = 0.0554 ± 0.0113 although there was no main effect of ASRS scores. We
(berries s−1)/patch number, p < 0.001; 5 s patches: OLS, found two-way effects of travel time and cumulative
βslope = 0.1269 ± 0.0174 (berries s−1)/patch number, p < 0.001). reward and of ASRS score and cumulative reward ( p < 5 ×
Of those participants who showed an effect of patch number 10−9 or less, Bonferroni-corrected). Finally, we also found a
 regression betas from patch-wise regression against time in patch
travel time
cumulative reward
ASRS score
travel time × cumulative reward
travel time × ASRS score
cumulative reward × ASRS score
Downloaded from https://royalsocietypublishing.org/ on 01 March 2024
travel time × cumulative reward ×
ASRS score
–0.1 0 0.1
Figure 4. Regression betas for fixed effects of travel time, cumulative reward, and ASRS scores and all interactions on time in patch (n = 10 353 patches). *: p < 0.0005
Bonferroni-corrected.
three-way interaction effect of travel time, cumulative reward
and ASRS score ( p < 0.0005, Bonferroni-corrected). Figure 4
depicts the regression betas for the covariates of interest.
We also examined trial-level effects of ASRS, testing our
prediction that participants who score highly on the ASRS
are more likely to choose to leave patches, all else being
equal. We performed a mixed-effects binomial regression of
all decisions to stay (= 0) or leave (= 1) a patch (n = 55 526
trials) against a random effect of patch number in session
and participant, fixed effects of patch residence time, travel
time, cumulative reward in patch and ASRS, including all
interactions of the fixed effect covariates, and a number
of nuisance covariates and their interactions (see §2d). All
fixed effect covariates were z-scored. Consistent with our pre-
dictions, all main fixed effects of covariates of interest ( p <
0.005 or less, Bonferroni-corrected) as well as all their two-
way interactions ( p < 5 × 10−5 or less, Bonferroni-corrected)
were significant. The three-way interaction of time in patch,
cumulative reward and ASRS score was also significant
( p < 5 × 10−7, Bonferroni-corrected), as was the four-way
interaction ( p < 5 × 10−8, Bonferroni-corrected). Figure 5
depicts the betas for the covariates of interest.
In the light of these significant interaction effects, we next
compared participants who screened positive on the ASRS
scale (n = 206) with those who screened negative (n = 251), the
clinically relevant comparison. A positive screen—a score of
14 or more—indicates that the individual exhibits symptoms
consistent with an ADHD diagnosis. Consistent with
our predictions, ASRS screen-positive participants stayed in
patches for significantly shorter durations than screen-negative
(Student’s t-test, p < 0.005, t (d.f. = 455) = −3.0085; mean
median time in patch for positive TiP+ = 19.3170 ± 0.9057 s;
TiP– = 23.3907 ± 0.9758 s). However, unexpectedly, they also
attained significantly higher reward rates than screen-negative
7
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Proc. R. Soc. B 291: 20222584
0.2 0.3 0.4 0.5 0.6 0.7 0.8
betas
participants (Student’s t-test, p < 0.00001, t (df = 455) = 4.5466;
mean reward rate for positive RR+ = 1.5896 ± 0.0356 points s–1;
RR– = 1.3608 ± 0.0350), contrary to our prediction. Though
ASRS screen-positive participants stayed less long in patches
than screen-negative participants, and earned higher reward
rates, they still tended to overharvest patches, staying signifi-
cantly longer than the optimal leave times predicted by the
MVT for both patch types (one-sample t-test against optimal
TiP; mean median TiP for 1 s patches = 17.2336 ± 1.0061 s,
p < 1 × 10−22, t (df = 204) = 11.1059; mean median TiP for 5 s
patches = 21.2876 ± 0.9211, p < 1 × 10−16, t (df = 204) = 9.0785).
Figure 6 illustrates the consistently shorter patch residence
times in both conditions for ADHD screen-positive compared
to screen-negative participants. Figure 7 illustrates the right-
shifted distribution of ADHD positive reward rates after
segregation by positive or negative ASRS screening. Cumulat-
ive rewards across sessions were significantly greater for
ADHD screen-positive than screen-negative participants
(Student’s t-test, p < 0.0001, t (df = 455) = 4.1000; mean cumulat-
ive reward for screen-positive C+ = 602.6460 ± 9.5222;
screen-negative C− = 521.6306 ± 10.0790).
We performed two sets of analyses to provide further
support for our conclusions. First, we ran the same
regressions and approximated the comparisons using OLS
with a simpler scoring scale for the ASRS screen: the sum
of the scores of the answers to each item on the questionnaire.
The results were almost identical to those reported above (see
electronic supplementary material). Second, 24 of our partici-
pants reported a previous diagnosis of ADHD (of those 24, 18
screened positive using the RiskSLIM scoring sheet [33]). We
again ran the same regressions and the analyses as above,
comparing those 24 participants with the 428 participants
who reported no previous ADHD diagnosis (5 participants
who did not answer that question were left out). The results
 regression betas from trial-wise binomial regression against leave decision 8

royalsocietypublishing.org/journal/rspb
time in patch
travel time *
cumulative reward *
ASRS score *
time in patch x travel time *
time in patch × cumulative reward *
travel time × cumulative reward *
time in patch × ASRS score *
travel time × ASRS score *
*

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cumulative reward × ASRS score
time in patch × travel time × *
cumulative reward
time in patch × travel time ×
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ASRS score
time in patch × cumulative reward ×
ASRS score
travel time × cumulative reward × *
ASRS score
time in patch × travel time ×
cumulative reward × ASRS score
*
–1.5 –1.0 –0.5 0 0.5 1.0 1.5 2.0 2.5 3.0
betas
Figure 5. Binomial regression betas for fixed effects of time in patch, travel time, cumulative reward and ASRS scores, and all interactions on stay or leave patch
decisions (n = 55 526 trials). *: p < 0.005 Bonferroni-corrected.

median patch residence times by ASRS screen outcomes new one. Participants made these decisions under short or
long travel times between patches, reflecting rich and poor
28
screen – environments. As predicted by the marginal value theorem
screen + (MVT), participants stayed longer in patches when travel
26
median total time in patch (s)

times were longer. Consistent with a range of evidence across

species [45,47,49,51], including humans [52,53], participants
24 stayed longer in patches than predicted by the MVT, potentially
reflecting gathering information about the environment [54], a
22 lack of competing demands due to predation or socializing, or
misrepresentation of rewards [55] or time [49]. The influence of
travel times on patch residence time resulted in significantly
20
different total rewards collected but not significantly different
reward rates between conditions.
18 We next tested for links between foraging decisions and
ADHD Self-Report Scale (ASRS) scores. Correcting for nui-
16 sance covariates, we found that interactions between ASRS
1 5
scores and cumulative reward, travel time and patch number
travel time (s)
in session predicted earlier patch leaving. A trial-level binomial
Figure 6. Patch residence times for ASRS screen-negative (blue) and screen- regression also uncovered significant interactions between
positive (red) participants for 1 s travel time (left) and 5 s travel time (right) ASRS and cumulative reward, patch number in session and
patches. Points are mean median total times in patch, error bars are ±1 s.e.m. time in patch. Comparing ASRS screen-positive with screen-
negative participants revealed shorter patch duration times
and higher reward rates for screen-positive individuals. We
were once again largely confirmatory (see electronic sup- followed-up and supported our findings with two sets of
plementary material), albeit noisier than the outcome of the further analyses, performing the same set of analyses using
first set of confirmatory analyses, perhaps due to so few the sum of the scores of the individual ASRS questions and
participants receiving previous ADHD diagnoses. comparing participants who had received a previous ADHD
diagnosis with those who had not.
There are several strengths and weaknesses to our design.
Our online design enabled gathering ADHD symptom self-
4. Discussion reports from many participants during the height of the
Foraging poses a fundamental challenge for humans and other COVID-19 pandemic. However, our findings relied on self-
organisms [12]. Here, we investigated how humans choose report without monitoring task engagement. Finally, although
between staying in a depleting resource patch or leaving for a the ASRS has been validated [33] and re-validated [40–44] and
 distributions of reward rates across all patches 9
0.14

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screen –
screen +
0.12

0.10

frequency 0.08

0.06

Proc. R. Soc. B 291: 20222584

0.04

0.02
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0.1

1.1

2.1
0
0.3

1.3

2.3
0.5

1.5

2.5
0.8

1.8

2.8
0.7

1.7

2.7
0.9

1.9

2.9
0.6

1.6

2.6
0.2

1.2

2.2
0.4

1.4

2.4
1.0

2.0

3.0
observed reward rate (pts s–1)
Figure 7. Distribution of patch reward rates across all patches for ASRS screen-negative (blue) and screen-positive (red) participants. Large peaks at 0 are from
participants departing patches immediately upon entry. Vertical dashed lines are optimal reward rates (see §2e): red: 1 s travel time optimal reward rate; black:
global optimal reward rate; green: 5 s travel time optimal reward rate.

self-report scales have been found to be informative for detect-
ing ADHD [56–58], clinical validation is still needed to confirm
the relationship between foraging behaviour and ADHD.
In our sample, 206 of 457 participants, or 45%, screened
positive for ADHD, approximately 10x higher than the rate
of ADHD in the adult population. Several factors may account
for this discrepancy. First, ADHD-screen positive participants
were not diagnosed with ADHD; only a clinical assessment
can confirm such a diagnosis. Second, we conducted our
study online using Prolific, which may have elevated rates of
positive screens, as was found using an older ADHD screen
in a previous mTurk sample [59]. Finally, our data collection
occurred during the COVID-19 pandemic, which negatively
impacted mental health for broad swaths of the public [60],
as confirmed by online self-reports [61]. Notably, the pandemic
is linked to a global increase in ADHD symptoms [62]. While
difficult to estimate, this context undoubtedly played a role
in the frequency of ADHD symptoms reported in our sample.
Higher reward rates in ADHD screen-positive participants
may reflect adaptive specialization in the capacity to focus.
Attention is known to be regulated in part by dopaminergic
receptors and transporters [63,64]. Animal models of ADHD
show dopaminergic deficits in the basal ganglia and prefrontal
cortex [65], and both the D2 [66] and the D4 [67] dopamine
receptors are associated with ADHD. These same receptors
are implicated in differences in nutrition and health in compari-
sons of nomadic and settled populations [27]; specifically, the
interaction between lifestyle and allele length for the D4 recep-
tor predicts healthier body mass. Longer alleles for the D4
receptor are a risk factor for ADHD [67]. Taken together,
allele length for the D4 receptor is a risk factor for ADHD
and correlates with better nutritive outcomes in nomadic
populations. Consistent with our reported findings, we specu-
late that ADHD serves as an adaptive specialization for
foraging [68–71], thus explaining its widespread prevalence
and continued persistence in the human population. Individ-
uals with ADHD may be more reward-seeking, consistent
with the role of dopamine in motivated behaviour [72] as
well as heightened impulsivity on delayed discounting tasks
[73]. Individuals with ADHD may also be more exploratory
as a result of increased noradrenergic drive originating in the
locus coeruleus and broadcast to prefrontal cortex [29]. Typical
individuals, by contrast, may be driven by a variety of motiv-
ations, including gathering more information about the
environment through persistent foraging.
Our task focuses on core explore–exploit foraging
decisions, a type of decision ubiquitous in the real world. For
example, the Aché people of Paraguay hunt in accordance
with the predictions of optimal foraging theory [74], as do
Nahua mushroom foragers [75], and human shopping behav-
iour is also consistent with optimal foraging predictions [76–
78]. However, there are key differences between naturalistic
foraging behaviour and patch leaving tasks like ours. Notably,
there are minor metabolic costs to our task, whereas naturalistic
foraging involves substantial metabolic costs associated with
leaving a patch and searching for a new one. In addition,
our task required participants to gather points—an abstract
resource that stands in contrast to basic alimentary or
other resources like calories. Nonetheless, animals and
humans show similar patterns of behaviour that accord with
foraging theory, whether foraging for resources in the real
world or foraging for food pellets or points on a computer in
the laboratory [22,50,55,79].
Our task was designed to understand how participants
make patch leaving decisions and whether participants with
different cognitive phenotypes exhibit variability in those
decisions. These decisions are characterized by tracking
resource intakes and deciding to replenish that resource. In
the real world, this decision would involve searching for a
new resource instead of merely resetting it. Unlike the real
 world, our task does not require search (cf. the distinction
between searching and harvesting in [80]). Nonetheless, by
focusing on patch leaving decisions, our task provides funda-
mental insights on foraging choices without the confounding
factors involved in searching for new patches.
ADHD-like cognitive phenotypes confer advantages or
disadvantages depending on the environment. Foragers with
ADHD-like traits may fare poorly in environments with mul-
tiple, learnable depletion rates (like the task used in [79]),
by leaving patches before gathering enough information to
learn the different patch types. In environments in which
exhausted patches are renewed, primates are known to track
renewal rates, such as the availability of fruit [81–83]. In these
environments, ADHD-type foragers may be likelier to visit
resources too early or too late due to noisier estimates of
depletion rates, resulting in fewer rewards. However, in
competitive environments where foragers must keep track of
other foragers, impulsively leaving patches could yield a
Downloaded from https://royalsocietypublishing.org/ on 01 March 2024
competitive advantage by enabling learning about competitors
and capturing newly renewed resources first.
Many decision contexts differ from foraging ones by pre-
senting agents with multiple options simultaneously rather
than single depleting resources. We hypothesize that individ-
uals with ADHD-type traits may struggle more in these
contexts, a proposal backed by existing research [84]. For
instance, a study by Frank et al. [85] used a reinforcement learn-
ing task to assess learning in neurotypical (healthy) individuals,
ADHD patients on medication and ADHD patients off medi-
cation. The participants had to learn which of two arbitrary
symbols was associated with more rewards by learning the
expected value of selecting each option. In their task, outcomes
were independent of previous choices, there was no depleting
resource and choices were between options instead of accepting
or rejecting a single offer. They found that patients off medi-
cation displayed lower choice accuracy and switched choices
more frequently after rewarding feedback than patients on
medication or healthy controls. This study illustrates how
in an environment with symbolic associations between rewards
and options, switching leads to maladaptive information
gathering to learn those associations, in contrast to a patchy
foraging environment like our task, where switching behaviour
may be adaptive.
As a second example, a study by Dekkers et al. [86] investi-
gated how ADHD participants and healthy controls made
choices between a more certain ‘safe’ option and a ‘risky’
option, each with explicitly verbally communicated expected
values. On some trials the safe option had the higher expected
value, whereas on others the risky option had a higher
expected value. ADHD participants less frequently selected
the option with the higher expected value. This decision con-
text involves a two-alternative forced choice where each
decision was independent and non-depleting, differing mark-
edly from the patch foraging task. Here, neither switching
nor staying was an intrinsically adaptive strategy, and optimal
decisions required information gathering and deployment (i.e.
in reading and computing the expected value of options).
Modern decision contexts often involve non-depleting, inde-
pendent choices, usually framed in symbolic terms. Such
tasks may pose specific challenges for individuals with
ADHD-type traits, whose decision-making mechanisms may
be better suited for foraging-like environments.
A burgeoning literature reveals links between foraging pro-
ficiency and attention to ADHD-like traits. Foraging for cryptic
prey types places attentional demands that lead to selection of 10
prey of a particular type [87], consistent with diet selection
royalsocietypublishing.org/journal/rspb
rules in optimal foraging theory [1]. In addition, a comparison
of feature search and conjunctive search revealed that search
type augments attentional demands and consequently fora-
ging costs, resulting in differences in give-up times [88], a key
foraging measure [89]. Finally, search paths are longer and
more variable in individuals with greater ADHD symptoms
[90]. Our findings build on this literature, suggesting
that heightened ADHD traits may confer certain foraging
advantages by lowering exploration thresholds.
The DMN, a distributed neural circuit with high resting-
state activity [28,29], shows elevated activity during task per-
formance in individuals with ADHD [30,31], potentially
Proc. R. Soc. B 291: 20222584
accounting for more frequent errors [29]. Neurons in posterior
cingulate cortex (PCC), a central node of the DMN, signal
decisions to explore [33] and the information driving these
decisions [18]. In addition, electrical microstimulation in PCC
promotes exploration of a non-default option [34]. ADHD is
associated with noradrenergic activity [28–30], and stimulant
medications like reboxetine [91] or atomoxetine [92] target
these systems. Notably, stimulation of the locus coeruleus—
the source of noradrenaline in the brain—provokes early
patch leaving in rats [32], and treatment with reboxetine in
humans induces earlier patch departures, higher reward rates
at patch leaves and more optimal foraging [50]. The locus coer-
uleus projects to a number of DMN nodes, including posterior
cingulate cortex [93,94]. Together, these findings suggest that
noradrenergic activity ‘tunes’ circuits regulating explore–
exploit decisions, resulting in advantages for ADHD-type indi-
viduals in foraging.
In conclusion, our virtual foraging task reveals that people
adaptively follow predictions of optimal foraging theory. They
adapted their foraging strategies to the local statistics of patches
and stayed longer in long travel time patches compared to
shorter travel times, as predicted by optimal foraging theory.
Nevertheless, people generally overharvested patches and con-
sequently did not conform to the optimal patch-leaving time
specified by foraging theory. In addition, we discovered that
participants that screened positive for ADHD more readily
abandoned patches and achieved higher reward rates than
did participants who screened negative. Given the over-staying
displayed by participants overall, those with elevated ASRS
scores made exploratory decisions that were more closely
aligned with the predictions of optimal foraging theory, and,
in this sense, behaved more optimally. The increased foraging
proficiency of participants with ADHD-like behaviour
observed here suggests that the prevalence and persistence of
ADHD in human populations may serve an adaptive function
in some environments.
Ethics. All methods were approved by the Internal Review Board of
the University of Pennsylvania (IRB #823436).
Data accessibility. All data are available at https://doi.org/10.5281/
zenodo.10178228 [95].
Supplementary material is available online [96].
Declaration of AI use. We have not used AI-assisted technologies in
creating this article.
Authors’ contributions. D.L.B.: data curation, formal analysis, software,
writing—original draft, writing—review and editing; V.U.L.: concep-
tualization, investigation, methodology, project administration,
supervision, writing—original draft, writing—review and editing;
F.P.: data curation, writing—review and editing; N.A.: investigation,
supervision; E.M.B.: writing—original draft, writing—review and
editing; A.R.: conceptualization, software; M.L.P.: conceptualization,
 funding acquisition, project administration, writing—original draft, Acknowledgements. The Wharton Behavioural Lab provided support 11
writing—review and editing. in setting up and running the study. We are grateful to Emily
All authors gave final approval for publication and agreed to be M. Orengo and Richard Lee for their assistance during this project.

held accountable for the work performed therein.
Conflict of interest declaration. We declare we have no competing interests.
Funding. The authors report no financial conflicts of interest or
disclosures.
royalsocietypublishing.org/journal/rspb
The research was supported by R37-MH109728, R01-MH108627,
R01-MH-118203, KA2019-105548, U01MH121260, UM1MH130981,
R56MH122819, R56AG071023 (all to M.L.P.), the Wharton Behavioural
Lab and the Wharton Dean’s Research Fund (to V.U.L. and M.L.P.).

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