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Yet, the mechanism linking CPS to peptidoglycan (PG) remains unclear. Here, we identified a strong negative genetic interaction between the genes encoding the putative capsule ligase CpsA and the WalK histidine kinase, a component of the WalRK two-component system regulating cell wall homeostasis. In the absence of\n                  <jats:italic>cpsA<\/jats:italic>\n                  , capsule polymers compete with wall teichoic acids for ligase activity to PG. This induces cell wall stress and is sensed by the WalRK system. Overexpression of the PG hydrolase\n                  <jats:italic>pcsB<\/jats:italic>\n                  or disruption of the PG-modifying enzymes\n                  <jats:italic>pgdA<\/jats:italic>\n                  and\n                  <jats:italic>oatA<\/jats:italic>\n                  restored growth of strains lacking\n                  <jats:italic>cpsA<\/jats:italic>\n                  and\n                  <jats:italic>walK<\/jats:italic>\n                  . Furthermore, CpsA overproduction compensates for the loss of other LytR-Cps2A-Psr (LCP) ligases, suggesting it can support capsule and wall teichoic acid syntheses. These findings support the model that LCP ligases are semi-redundant, although they may install secondary polymers on a different residue of PG. This work also suggests that WalRK signaling compensates for reduced capsule and WTA attachment by positively regulating PG hydrolases.\n                <\/jats:p>\n                <jats:sec>\n                  <jats:title>SIGNIFICANT STATEMENT<\/jats:title>\n                  <jats:p>\n                    <jats:italic>Streptococcus pneumoniae<\/jats:italic>\n                    causes approximately half a million deaths annually. A powerful public health tool for controlling pneumococcal infections is vaccination against the protective capsule. Yet, the mechanisms by which the capsule layer attaches to the underlying cell wall remain poorly defined. This study shows that the conserved capsule gene CpsA is not strictly required for capsule attachment but instead works together with other LytRIZCpsAIZPsr (LCP) ligases. However, it requires the essential WalRK signaling system to maintain cell envelope integrity. Defects in LCP activity are alleviated by WalRKIZdriven upregulation of peptidoglycan hydrolases, overexpression of PcsB, or inactivating peptidoglycan modifications that limit hydrolysis. These findings reveal coordination among flux to capsule synthesis, secondary wall polymer attachment, and cell wall remodeling.\n                  <\/jats:p>\n                <\/jats:sec>\n                <jats:sec>\n                  <jats:title>TEASER<\/jats:title>\n                  <jats:p>The WalRK two-component system responds to cell envelope stress caused by reduced LCP ligase activity.<\/jats:p>\n                <\/jats:sec>","DOI":"10.64898\/2026.03.25.714115","type":"posted-content","created":{"date-parts":[[2026,3,27]],"date-time":"2026-03-27T04:56:11Z","timestamp":1774587371000},"source":"Crossref","is-referenced-by-count":0,"title":["The WalRK two-component system in\n                  <i>Streptococcus pneumoniae<\/i>\n                  ensures robustness of secondary wall polymer attachment"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0260-2446","authenticated-orcid":false,"given":"Justin J.","family":"Zik","sequence":"first","affiliation":[{"name":"Infectious Diseases Translational Research Programme and Department of Microbiology and Immunology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zeyu","family":"Fu","sequence":"additional","affiliation":[{"name":"Infectious Diseases Translational Research Programme and Department of Microbiology and Immunology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Morgan N.","family":"Price","sequence":"additional","affiliation":[{"name":"Environmental Genomics and Systems Biology Division, Lawrence Berkeley National Laboratory, Berkeley, CA 94720, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yujie","family":"Li","sequence":"additional","affiliation":[{"name":"School of Chemistry, Chemical Engineering and Biotechnology (CCEB), Nanyang Technological University (NTU), Singapore"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuan","family":"Qiao","sequence":"additional","affiliation":[{"name":"School of Chemistry, Chemical Engineering and Biotechnology (CCEB), Nanyang Technological University (NTU), Singapore"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4999-2931","authenticated-orcid":false,"given":"Adam P.","family":"Arkin","sequence":"additional","affiliation":[{"name":"Environmental Genomics and Systems Biology Division, Lawrence Berkeley National Laboratory, Berkeley, CA 94720, USA"},{"name":"Department of Bioengineering, University of California, Berkeley, Berkeley, CA 94720, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Adam M.","family":"Deutschbauer","sequence":"additional","affiliation":[{"name":"Environmental Genomics and Systems Biology Division, Lawrence Berkeley National Laboratory, Berkeley, CA 94720, USA"},{"name":"Department of Plant and Microbial Biology, University of California, Berkeley, CA 94720, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Josue","family":"Flores-Kim","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Molecular Biotechnology, UMass Chan Medical School, Worcester, MA, 01605, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9047-0310","authenticated-orcid":false,"given":"Lok-To","family":"Sham","sequence":"additional","affiliation":[{"name":"Infectious Diseases Translational Research Programme and Department of Microbiology and Immunology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"54368","reference":[{"key":"2026032910431177000_2026.03.25.714115v1.1","doi-asserted-by":"publisher","DOI":"10.1038\/s41579-018-0001-8"},{"key":"2026032910431177000_2026.03.25.714115v1.2","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(22)02185-7"},{"key":"2026032910431177000_2026.03.25.714115v1.3","doi-asserted-by":"publisher","DOI":"10.1016\/S1473-3099(24)00176-2"},{"key":"2026032910431177000_2026.03.25.714115v1.4","doi-asserted-by":"crossref","first-page":"e744","DOI":"10.1016\/S2214-109X(18)30247-X","article-title":"Burden of Streptococcus pneumoniae and Haemophilus influenzae type b disease in children in the era of conjugate vaccines: global, regional, and national estimates for 2000-15","volume":"6","year":"2018","journal-title":"Lancet Glob Health"},{"key":"2026032910431177000_2026.03.25.714115v1.5","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(24)01867-1\/ATTACHMENT\/4F9FA746-A486-4BC0-BF71-075D4D1D6D56\/MMC3.PDF"},{"key":"2026032910431177000_2026.03.25.714115v1.6","unstructured":"Active Bacterial Core Surveillance (ABCs) Report Emerging Infections Program Network=: Streptococcus pneumoniae , 2018. 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