ZIKA VIRUS IMPAIRS NEUROBEHAVIORAL DEVELOPMENT AND INDUCES OXIDATIVE STRESS LINKED TO BLOOD–BRAIN BARRIER DISRUPTION IN A RAT MODEL OF CONGENITAL INFECTION

Аннотация

Zika virus (ZIKV) is an arbovirus belonging to the Flaviviridae family and the Flavivirus genus, which caused a significant epidemic in the Americas in 2015 (Song et al., 2017). Gestational ZIKV infection was soon linked to neurodevelopmental complications in fetuses, leading to various adverse outcomes such as intrauterine growth restriction, fetal death, miscarriage, stillbirth, and ocular abnormalities (Alvarado & Schwartz, 2017; Coyne & Lazear, 2016). Among these, microcephaly emerged as the most prominent clinical manifestation and is now well-established as a consequence of ZIKV vertical transmission (Wang & Ling, 2016; Li et al., 2016). While microcephaly was initially considered the primary neurological outcome of congenital ZIKV infection, recent studies have demonstrated that children affected by ZIKV in utero, even without microcephaly, may still exhibit neurodevelopmental delays (Sobral da Silva et al., 2021). Furthermore, growing evidence suggests that ZIKV-induced pathogenesis extends beyond the immediate postnatal period, with potential long-term effects on neurological development.

Тип источника: Конференции
Годы охвата с 2022
inLibrary
Google Scholar
Выпуск:
93-97
22

Скачивания

Данные скачивания пока недоступны.
Поделиться
Kenjayev , B. (2025). ZIKA VIRUS IMPAIRS NEUROBEHAVIORAL DEVELOPMENT AND INDUCES OXIDATIVE STRESS LINKED TO BLOOD–BRAIN BARRIER DISRUPTION IN A RAT MODEL OF CONGENITAL INFECTION. Естественные науки в современном мире: теоретические и практические исследования, 4(1), 93–97. извлечено от https://inlibrary.uz/index.php/zdtf/article/view/65026
Crossref
Сrossref
Scopus
Scopus

Аннотация

Zika virus (ZIKV) is an arbovirus belonging to the Flaviviridae family and the Flavivirus genus, which caused a significant epidemic in the Americas in 2015 (Song et al., 2017). Gestational ZIKV infection was soon linked to neurodevelopmental complications in fetuses, leading to various adverse outcomes such as intrauterine growth restriction, fetal death, miscarriage, stillbirth, and ocular abnormalities (Alvarado & Schwartz, 2017; Coyne & Lazear, 2016). Among these, microcephaly emerged as the most prominent clinical manifestation and is now well-established as a consequence of ZIKV vertical transmission (Wang & Ling, 2016; Li et al., 2016). While microcephaly was initially considered the primary neurological outcome of congenital ZIKV infection, recent studies have demonstrated that children affected by ZIKV in utero, even without microcephaly, may still exhibit neurodevelopmental delays (Sobral da Silva et al., 2021). Furthermore, growing evidence suggests that ZIKV-induced pathogenesis extends beyond the immediate postnatal period, with potential long-term effects on neurological development.


background image

93

ZIKA VIRUS IMPAIRS NEUROBEHAVIORAL DEVELOPMENT AND INDUCES

OXIDATIVE STRESS LINKED TO BLOOD–BRAIN BARRIER DISRUPTION IN A

RAT MODEL OF CONGENITAL INFECTION

Kenjayev Baxtiyor

Assistant lecturer at the Alfraganus University

Email address: ismatovbaxtiyor.67890.@gmail.com

ORCID ID: 0000-0003-3259-8492

https://doi.org/10.5281/zenodo.14783854

Introduction

Zika virus (ZIKV) is an arbovirus belonging to the

Flaviviridae

family and the

Flavivirus

genus, which caused a significant epidemic in the Americas in 2015 (Song et al., 2017).
Gestational ZIKV infection was soon linked to neurodevelopmental complications in fetuses,
leading to various adverse outcomes such as intrauterine growth restriction, fetal death,
miscarriage, stillbirth, and ocular abnormalities (Alvarado & Schwartz, 2017; Coyne & Lazear,
2016). Among these, microcephaly emerged as the most prominent clinical manifestation and
is now well-established as a consequence of ZIKV vertical transmission (Wang & Ling, 2016;
Li et al., 2016). While microcephaly was initially considered the primary neurological outcome
of congenital ZIKV infection, recent studies have demonstrated that children affected by ZIKV
in utero, even without microcephaly, may still exhibit neurodevelopmental delays (Sobral da
Silva et al., 2021). Furthermore, growing evidence suggests that ZIKV-induced pathogenesis
extends beyond the immediate postnatal period, with potential long-term effects on
neurological development.

Several models have been developed to explore the pathophysiology of ZIKV infection

and its impact on the nervous system.

In vitro

studies have been instrumental in elucidating

the cellular and molecular mechanisms underlying ZIKV-induced neurological damage. For
instance, studies using U87-MG (human glioblastoma) and HepG2 (human liver carcinoma)
cell lines have revealed that ZIKV infection leads to increased reactive oxygen species (ROS),
lipid peroxidation, and protein carbonylation, alongside a decrease in antioxidant enzyme
activity, including superoxide dismutase and catalase (Almeida et al., 2020). Similarly, human
neural progenitor cells (hNPCs) infected with ZIKV exhibited cell cycle dysregulation and
increased caspase-3 activation, ultimately leading to cell death (Tang et al., 2016). In radial
glial cells (RGCs), ZIKV infection disrupts mitosis and structural organization and leads to the
sequestration of phosphorylated TBK1 during mitosis (Onorati et al., 2016). Given the crucial
role of RGCs in cortical cell migration, these findings help explain the cortical thinning
observed in ZIKV-infected models.

Another significant consequence of ZIKV infection involves damage to the blood–brain

barrier (BBB), a critical structure that regulates the exchange between systemic circulation
and the central nervous system (Clé et al., 2020). Studies conducted

in vitro

and

in vivo

have

demonstrated that ZIKV infection disrupts BBB integrity (Leda et al., 2019). However, it
remains unclear whether these alterations are transient or contribute to long-term
neuropathological outcomes. Additionally, the full spectrum of BBB-related damage induced
by ZIKV infection has yet to be fully elucidated.

Regarding

in vivo

studies, various rodent models have been employed to investigate

ZIKV pathogenicity, fetal infection, and vertical transmission. Since wild-type (WT) rodents


background image

94

exhibit resistance to ZIKV due to their robust interferon (IFN) response, many studies have
relied on IFN-deficient animal models to assess infection-related outcomes (Kublin &
Whitney, 2018). Immunocompromised mice, such as A129 and AG129 strains, have been
instrumental in understanding the impact of vertical transmission (Vue & Tang, 2021).
Additionally, alternative models have explored different routes of infection, including direct
intrauterine exposure to the developing rodent brain (Li et al., 2016) and postnatal infection
shortly after birth (Lazear et al., 2016; Miner et al., 2016). Given the need to better understand
immune responses, long-term consequences, and potential therapeutic interventions, further
research involving immunocompetent animal models is essential (Morrison & Diamond,
2017).


Methods
Animal Procedures

This study utilized pregnant female Wistar rats, approximately two months old,

obtained from the Animal Reproduction and Experimental Center at the Federal University of
Rio Grande do Sul, Brazil. The animals were housed in individually ventilated cages (IVC)
under a 12-hour light/dark cycle, with a controlled temperature of 21 ± 2°C, and provided
with unrestricted access to food and water. All experimental procedures adhered to the
ethical guidelines established by the National Council for Animal Experimentation of Brazil
and complied with Brazilian legislation for the scientific use of animals (Law 11.794/08).
Additionally, procedures followed the

Guide for the Care and Use of Laboratory Animals

as

outlined by the National Research Council (USA, 2011).

Ethical approval for this study was granted by the Ethics Committee of the Federal

University of Rio Grande do Sul (approval number 33452/2017). All procedures were
conducted in a Biosafety Level 2 (BSL-2) laboratory under Animal Biosafety Level 2 (ABSL-2)
conditions, in accordance with the guidelines of the Centers for Disease Control and
Prevention (CDC) for laboratory work involving Zika virus. Virus handling and animal-related
procedures were performed within a Class II Biological Safety Cabinet (BSC) (Tecniplast® BS
60 class II, Buguggiate, Italy) to ensure biosafety compliance.

On embryonic day 9 (E9), pregnant females were randomly assigned to two groups. Five

females received an intraperitoneal injection of 500 µL containing

1 × 10⁶

plaque-forming

units per milliliter (PFU/mL) of Zika virus isolated in Brazil (

ZIKV_BR

). The control group

consisted of six females that received a 500 µL intraperitoneal injection of sterile diluted
medium.

Neurological Reflexes

Twenty-four hours after birth, litter standardization (5–8 pups per litter) and div

measurements were conducted (Table 1). To assess neurodevelopmental impairments
associated with gestational Zika virus infection, a series of neurological reflex evaluations
were performed. These assessments began on postnatal day (PND) 3 and were conducted
every three days until PND 21. Observations included physical characteristics such as div
weight, eye opening, and incisor tooth eruption (Lubics et al., 2005).

The following reflexes and motor responses were evaluated:

Righting Reflex:

Each pup was placed on its back, and the time taken to return to a

prone position with all four paws in contact with the surface was recorded.


background image

95

Negative Geotaxis:

Pups were positioned on an inclined platform (45°) with their heads

facing downward. The day when they were able to turn their heads upward and climb the
platform was noted, with a maximum time limit of 30 seconds to complete the task.

Limb Placing:

The posterior region of the forepaw and hind paw was gently pressed

against a surface, and the day when the pup successfully placed its paws onto the surface was
recorded.

Limb Grasp:

The back of the forelimb paws was lightly touched against a stem, and the

day when the pup grasped the stem was noted.

Cliff Aversion:

Each pup was placed with its head near the edge of an elevated platform,

and the time taken to turn its head away from the edge was measured. Additionally, the first
day this behavior was observed was recorded.

Gait Assessment:

Pups were placed at the center of a 30 cm-diameter circle, and the

time taken to exit the circle was recorded, along with the first day they exhibited locomotor
movement.

Results
Day of Appearance of Neurological Reflexes

To assess the emergence of neurological reflexes, a

t-test

was conducted, comparing

male ZKV (Zika virus-infected) with male CT (control) and female ZKV with female CT. A
significant delay in the onset of specific neurological reflexes was observed in male ZKV
compared to male CT in the following parameters:

Incisor tooth eruption

(t(28) = 5.82; P < 0.05)

Forelimb placing (right)

(t(28) = 4.58; P < 0.05)

Hind limb placing (right)

(t(28) = 5.15; P < 0.05)

Hind limb placing (left)

(t(28) = 3.35; P < 0.05)

Forelimb grasp (right)

(t(28) = 2.15; P < 0.05)

No significant differences were detected in the timing of

eye opening, negative

geotaxis, forelimb placing (left), forelimb grasp (left), gait, aversion to fall, righting
reflex, or olfactory behavior

.

For females, a similar analysis revealed delayed reflex development in the following:

Hind limb placing (right)

(t(13) = 3.10; P < 0.05)

Hind limb placing (left)

(t(13) = 4.56; P < 0.05)

Righting reflex

(t(28) = 2.28; P < 0.05)

No significant differences were found in other assessed reflexes.
Overall, these findings indicate that gestational Zika virus infection resulted in delays in

the appearance of certain neurological reflexes in both male and female offspring. These
delays may serve as early indicators of potential long-term neurological alterations.

Data are presented as

mean ± SD

.

(ZKV)

refers to offspring from Zika virus-infected

mothers, while

(CT)

represents control animals. Sample sizes: Female CT

(n = 8)

, Female ZKV

(n = 7)

, Male CT

(n = 15)

, Male ZKV

(n = 15)

. Asterisks (*) indicate significant differences

compared to the control group of the same sex (Student’s t-test, p < 0.05).

Discussion

In this study, we expanded the understanding of

congenital Zika virus (ZIKV) syndrome

by developing a rat model of congenital ZIKV infection to investigate

neurodevelopmental


background image

96

impairments and brain tissue disturbances

in Wistar rats shortly after birth. The infection was

induced on

embryonic day 9 (E9)

,

a crucial stage for rodent neurodevelopment (Semple et al.,

2013). Pregnant rats received an

intraperitoneal (i.p.) injection

of ZIKV, and viral presence

was detected in maternal blood as early as

six hours post-infection

.

Additionally, viable virus

was found in the

placenta, spleen, and fetuses

within 24 hours post-inoculation. Notably

,

infected females did not exhibit sickness behavior

despite their offspring showing

neurological impairments, which aligns with previous studies (Sherer et al., 2019). This
finding is particularly relevant, given that

asymptomatic ZIKV circulation has been reported

among pregnant women

in northeastern Brazil (Branco et al., 2021). Furthermore, even

asymptomatic maternal ZIKV infection

has been associated with

neurodevelopmental

impairments in offspring

(Shapiro-Mendoza et al., 2017). Consistently, our study identified

significant neurobehavioral deficiencies

in infected pups, suggesting potential

long-term

cognitive and motor disturbances

.

On

postnatal day 22 (PND 22)

,

blood–brain barrier (BBB)

integrity deficits

in the hippocampus and

altered oxidative status

in the hippocampus and

cortex were observed, highlighting the potential for lasting structural and functional brain
damage

.

Conclusion

This study sought to deepen the understanding of the long-term effects of gestational

Zika virus (ZIKV) infection by developing an immunocompetent rat model and identifying
neurobehavioral markers predictive of future disabilities. Our findings reveal that, even in the
absence of brain morphometric changes or a microcephaly-like phenotype, infected offspring
exhibited significant neurodevelopmental impairments. These deficits were associated with
hippocampal and cortical blood–brain barrier (BBB) disruption

and oxidative stress

imbalance observed 22 days postnatally

.

Together, these results provide strong evidence that the impact of gestational ZIKV

infection extends beyond the neonatal period, with potential long-term consequences for
brain function and development

.

Our study contributes to existing knowledge by highlighting

the need for improved prenatal diagnostics, early interventions, and extended follow-up
studies to further elucidate the mechanisms underlying congenital ZIKV syndrome.
Furthermore, the development of a reliable congenital infection model not only enhances our
understanding of disease pathology but also offers new opportunities for advancing
therapeutic strategies

.

References:

1.

J.B. Alimonti, M. Ribecco-Lutkiewicz, C. Sodja, A. Jezierski, D.B. Stanimirovic, Q. Liu, A.S.

Haqqani, W. Conlan, M. Bani-Yaghoub “

Zika virus crosses an in vitro human blood brain barrier

model

Fluids Barrier” CNS

, 15(1) (2018), 10.1186/S12987-018-0100-Y

2.

L.T. Almeida, A.C. Ferraz, C.C. da Silva Caetano, M.B. da Silva Menegatto, A.C. dos Santos

Pereira Andrade, R.L.S. Lima, F.C. Camini, S.H. Pereira, K.Y. da Silva Pereira, B. de Mello Silva,
L.O. Perucci, A. Talvani, J.C. de Magalhães, C.L. de Brito Magalhães

Zika virus induces oxidative

stress and decreases antioxidant enzyme activities in vitro and in vivo

Virus Res.

, 286 (2020), p.

198084
3.

M.G. Alvarado, D.A. Schwartz “

Zika Virus Infection in Pregnancy, Microcephaly, and

Maternal and Fetal Health: What We Think, What We Know, and What We Think We Know

Arch.


background image

97

Pathol. Lab. Med.

,” 141(1) (2017), pp. 26-32, 10.5858/arpa.2016-0382-RA

4.

M.F.V.V. Aragao, A.C. Holanda, A.M. Brainer-Lima, N.C.L. Petribu, M. Castillo, V. van der

Linden, S.C. Serpa, A.G. Tenório, P.T.C. Travassos, M.T. Cordeiro, C. Sarteschi, M.M. Valenca, A.
Costello

Nonmicrocephalic Infants with Congenital Zika Syndrome Suspected Only after

Neuroimaging Evaluation Compared with Those with Microcephaly at Birth and Postnatally:
How Large Is the Zika Virus “Iceberg”?

AJNR Am J Neuroradiol.

, 38(7) (2017), pp. 1427-1434

5.

G. Barisano, A. Montagne, K. Kisler, J.A. Schneider, J.M. Wardlaw, B.V. Zlokovic

Blood–brain barrier link to human cognitive impairment and Alzheimer’s disease

Nat. Cardiovasc. Res.

, 1(2) (2022), 108–115,

https://doi.org/10.1038/s44161-021-00014-4

6.

R.C.C. Branco, P. Brasil, J.M.G. Araújo, F.O. Cardoso, Z.S. Batista, V.M.S. Leitão, M.A.C.N. da

Silva, L.O. de Castro, J.G. Valverde, S.M.B. Jeronimo, J.A. Lima, R. Ribeiro da Silva, M.D.C.L.
Barbosa, L.M.O. Brito, M.A.P. Xavier, M.D.D.S.B. Nascimento

Evidence of Zika virus circulation in

asymptomatic pregnant women in Northeast, Brazil

PLoS Negl. Trop. Dis.

, 15(6) (2021),

e0009412,

https://doi.org/10.1371/journal.pntd.0009412

7.

G.J. Burton, E. Jauniaux “

Oxidative stress

Best Pract. Res. Clin. Obstet. Gynaecol.

”25(3)

(2011), p. 287, 10.1016/J.BPOBGYN.2010.10.016
8.

G. Calvet, R.S. Aguiar, A.S. Melo, S.A. Sampaio, I. de Filippis, A. Fabri, E.S. Araujo, P.C. de

Sequeira, M.C. de Mendonca, L. de Oliveira, D.A. Tschoeke, C.G. Schrago, F.L. Thompson, P.
Brasil,

F.B.

dos

Santos,

R.M.

Nogueira,

A.

Tanuri,

A.M.

de

Filippis

“Detection and sequencing of Zika virus from amniotic fluid of fetuses with microcephaly in
Brazil: a case study

Lancet Infect. Dis.

,” 16(6) (2016), 653–660,

https://doi.org/S1473-

3099(16)00095-5

9.

Ibroximovna, M. S. (2024). FACTORS OF DEVELOPING OF INTERCULTURAL

COMMUNICATION COMPETENCE IN TEACHING ENGLISH TO CADETS OF MILITARY
UNIVERSITY. Лучшие интеллектуальные исследования, 15(1), 159-163.
10.

Musayeva, S. I. (2024, May). DEVELOPMENT OF INTERCULTURAL COMMUNICATION

COMPETENCE OF CADETS USING INTERACTIVE METHODS. In Proceedings of International
Conference on Scientific Research in Natural and Social Sciences (Vol. 3, No. 5, pp. 276-284).
11.

T.F. Cardoso, R.S. dos Santos, R.M. Corrêa, J.V. Campos, R.D.B. Silva, C.C. Tobias, A. Prata-

Barbosa, A.J.L.A. da Cunha, H.C. Ferreira “

Congenital Zika infection: neurology can occur

without microcephaly Arch. Dis. Child.

, 104(2) (2019), pp. 199-200, 10.1136/ARCHDISCHILD-

2018-314782
12.

M. Clé, C. Desmetz, J. Barthelemy, M.-F. Martin, O. Constant, G. Maarifi, V. Foulongne, K.

Bolloré, Y. Glasson, F. De Bock, M. Blaquiere, L. Dehouck, N. Pirot, E. Tuaillon, S. Nisole, F.
Najioullah, P. Van de Perre, A. Cabié, N. Marchi, F. Gosselet, Y. Simonin, S. Salinas, M.S.
Diamond “

Zika Virus Infection Promotes Local Inflammation, Cell Adhesion Molecule

Upregulation, and Leukocyte Recruitment at the Blood-Brain Barrier

MBio

,” 11(4) (2020),

10.1128/MBIO.01183-20

Библиографические ссылки

J.B. Alimonti, M. Ribecco-Lutkiewicz, C. Sodja, A. Jezierski, D.B. Stanimirovic, Q. Liu, A.S. Haqqani, W. Conlan, M. Bani-Yaghoub “Zika virus crosses an in vitro human blood brain barrier model Fluids Barrier” CNS, 15(1) (2018), 10.1186/S12987-018-0100-Y

L.T. Almeida, A.C. Ferraz, C.C. da Silva Caetano, M.B. da Silva Menegatto, A.C. dos Santos Pereira Andrade, R.L.S. Lima, F.C. Camini, S.H. Pereira, K.Y. da Silva Pereira, B. de Mello Silva, L.O. Perucci, A. Talvani, J.C. de Magalhães, C.L. de Brito Magalhães Zika virus induces oxidative stress and decreases antioxidant enzyme activities in vitro and in vivo Virus Res., 286 (2020), p. 198084

M.G. Alvarado, D.A. Schwartz “Zika Virus Infection in Pregnancy, Microcephaly, and Maternal and Fetal Health: What We Think, What We Know, and What We Think We Know Arch. Pathol. Lab. Med.,” 141(1) (2017), pp. 26-32, 10.5858/arpa.2016-0382-RA

M.F.V.V. Aragao, A.C. Holanda, A.M. Brainer-Lima, N.C.L. Petribu, M. Castillo, V. van der Linden, S.C. Serpa, A.G. Tenório, P.T.C. Travassos, M.T. Cordeiro, C. Sarteschi, M.M. Valenca, A. Costello Nonmicrocephalic Infants with Congenital Zika Syndrome Suspected Only after Neuroimaging Evaluation Compared with Those with Microcephaly at Birth and Postnatally: How Large Is the Zika Virus “Iceberg”? AJNR Am J Neuroradiol., 38(7) (2017), pp. 1427-1434

G. Barisano, A. Montagne, K. Kisler, J.A. Schneider, J.M. Wardlaw, B.V. Zlokovic

Blood–brain barrier link to human cognitive impairment and Alzheimer’s disease

Nat. Cardiovasc. Res., 1(2) (2022), 108–115, https://doi.org/10.1038/s44161-021-00014-4

R.C.C. Branco, P. Brasil, J.M.G. Araújo, F.O. Cardoso, Z.S. Batista, V.M.S. Leitão, M.A.C.N. da Silva, L.O. de Castro, J.G. Valverde, S.M.B. Jeronimo, J.A. Lima, R. Ribeiro da Silva, M.D.C.L. Barbosa, L.M.O. Brito, M.A.P. Xavier, M.D.D.S.B. Nascimento Evidence of Zika virus circulation in asymptomatic pregnant women in Northeast, Brazil PLoS Negl. Trop. Dis., 15(6) (2021), e0009412, https://doi.org/10.1371/journal.pntd.0009412

G.J. Burton, E. Jauniaux “Oxidative stress Best Pract. Res. Clin. Obstet. Gynaecol.”25(3) (2011), p. 287, 10.1016/J.BPOBGYN.2010.10.016

G. Calvet, R.S. Aguiar, A.S. Melo, S.A. Sampaio, I. de Filippis, A. Fabri, E.S. Araujo, P.C. de Sequeira, M.C. de Mendonca, L. de Oliveira, D.A. Tschoeke, C.G. Schrago, F.L. Thompson, P. Brasil, F.B. dos Santos, R.M. Nogueira, A. Tanuri, A.M. de Filippis

“Detection and sequencing of Zika virus from amniotic fluid of fetuses with microcephaly in Brazil: a case study Lancet Infect. Dis.,” 16(6) (2016), 653–660, https://doi.org/S1473-3099(16)00095-5

Ibroximovna, M. S. (2024). FACTORS OF DEVELOPING OF INTERCULTURAL COMMUNICATION COMPETENCE IN TEACHING ENGLISH TO CADETS OF MILITARY UNIVERSITY. Лучшие интеллектуальные исследования, 15(1), 159-163.

Musayeva, S. I. (2024, May). DEVELOPMENT OF INTERCULTURAL COMMUNICATION COMPETENCE OF CADETS USING INTERACTIVE METHODS. In Proceedings of International Conference on Scientific Research in Natural and Social Sciences (Vol. 3, No. 5, pp. 276-284).

T.F. Cardoso, R.S. dos Santos, R.M. Corrêa, J.V. Campos, R.D.B. Silva, C.C. Tobias, A. Prata-Barbosa, A.J.L.A. da Cunha, H.C. Ferreira “Congenital Zika infection: neurology can occur without microcephaly Arch. Dis. Child., 104(2) (2019), pp. 199-200, 10.1136/ARCHDISCHILD-2018-314782

M. Clé, C. Desmetz, J. Barthelemy, M.-F. Martin, O. Constant, G. Maarifi, V. Foulongne, K. Bolloré, Y. Glasson, F. De Bock, M. Blaquiere, L. Dehouck, N. Pirot, E. Tuaillon, S. Nisole, F. Najioullah, P. Van de Perre, A. Cabié, N. Marchi, F. Gosselet, Y. Simonin, S. Salinas, M.S. Diamond “Zika Virus Infection Promotes Local Inflammation, Cell Adhesion Molecule Upregulation, and Leukocyte Recruitment at the Blood-Brain Barrier MBio,” 11(4) (2020), 10.1128/MBIO.01183-20