the angular momentum polarization present in
the initial HD molecules has been lost. For both
H-SARP and V-SARP prepared HD, the scattered
molecules have not preserved the initial molecular
alignment. Although this behavior contradicts the
well-known Dm = 0 selection rule for collisions
(39–41), this is to be expected for molecular scattering where the depth of the anisotropic potential is
substantially greater than the collision energy.
As such, the observed depolarization provides
clear evidence of anisotropic interactions that
strongly couple the internal and orbital angular
momenta. These strong forces suggest that a
transient van der Waals complex may be present
at the collision energies of this experiment.
We can make an order-of-magnitude estimate
of the total collision cross section using a procedure that is described in the supplementary
materials. For the initial HD molecular state
prepared by V-SARP, we find an integral cross
section of ~42 Å2, whereas for the state prepared by H-SARP, it is reduced by a factor of 3
to ~14 Å2. Our estimated cross section seems to
agree with a prior theoretical prediction (30)
within an order of magnitude, suggesting again
the presence of a van der Waals resonance as
described in that reference.
Without both HD state preparation using
SARP and reduction in the number of involved
partial waves due to the cold collisions present
in the coexpanded molecular beam of mixed
gases, the remarkable stereodynamic information
revealed in these experiments would have been
obscured. This study demonstrates conclusively
the importance for collision dynamics of the preparation of initial quantum state of the entire system, including all possible degrees of freedom.
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This work has been supported by the U.S. Army Research
Office under Army Research Office (ARO) grant no. W911NF-16-1-
1061 and Multidisciplinary University Research Initiatives (MURI)
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Materials and Methods
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6 July 2017; accepted 8 September 2017
Ectopic colonization of oral bacteria
in the intestine drives TH1 cell
induction and inflammation
Koji Atarashi,1,2 Wataru Suda,1,3,4 Chengwei Luo,5,6 Takaaki Kawaguchi,1,2 Iori Motoo,2
Seiko Narushima,2 Yuya Kiguchi,3 Keiko Yasuma,1 Eiichiro Watanabe,2
Takeshi Tanoue,1,2 Christoph A. Thaiss,7 Mayuko Sato,8 Kiminori Toyooka,8
Heba S. Said,4,9 Hirokazu Yamagami,10 Scott A. Rice,11 Dirk Gevers,5
Ryan C. Johnson,12 Julia A. Segre,12 Kong Chen,13 Jay K. Kolls,13 Eran Elinav,7
Hidetoshi Morita,14 Ramnik J. Xavier,5,6 Masahira Hattori,3,4 Kenya Honda1,2*
Intestinal colonization by bacteria of oral origin has been correlated with several negative
health outcomes, including inflammatory bowel disease. However, a causal role of oral
bacteria ectopically colonizing the intestine remains unclear. Using gnotobiotic techniques,
we show that strains of Klebsiella spp. isolated from the salivary microbiota are strong
inducers of T helper 1 (TH1) cells when they colonize in the gut. These Klebsiella strains are
resistant to multiple antibiotics, tend to colonize when the intestinal microbiota is
dysbiotic, and elicit a severe gut inflammation in the context of a genetically susceptible
host. Our findings suggest that the oral cavity may serve as a reservoir for potential
intestinal pathobionts that can exacerbate intestinal disease.
The average person generates and ingests ~1.5 liters of saliva per day, containing an enormous number of oral-resident bacteria (1, 2). Ingested oral bacteria poorly colo- nize the healthy intestine (3); however, increased levels of microbes of oral origin have been
reported in the gut microbiota of patients with
several diseases, including inflammatory bowel
disease (IBD) (4), HIV infection (5, 6), liver cirrhosis (7, 8), and colon cancer (9). For instance, the
presence of oral bacteria such as Veillonellaceae
and Fusobacteriaceae in the intestinal mucosal
microbiota strongly correlates with disease status in Crohn’s disease (CD) (4). Mining of our in-house data sets of 16S ribosomal RNA (rRNA)
gene sequences revealed that several bacterial
taxa—including species belonging to Rothia,
Streptococcus, Neisseria, Prevotella, and Gemella
(table S1A), all of which are aerotolerant and
typically members of the oral microbiota—were
significantly more abundant in the fecal microbiota of patients with ulcerative colitis (UC), primary sclerosing cholangitis (PSC), gastroesophageal
reflux disease (GERD) being treated by long-term
proton pump inhibitor therapy, and alcoholism,
compared with that of healthy controls (Fig. 1A
and table S1B). Thus, we hypothesized that a
subset of oral microbiota may ectopically colonize
and persist in the intestine under certain circumstances to aberrantly activate the intestinal