articles of association presentation

• Search Search Please fill out this field.

What Are Articles of Association?

Understanding articles of association, small business example of articles of association, the bottom line.

• Business Essentials

Articles of Association Definition and Example in Small Business

James Chen, CMT is an expert trader, investment adviser, and global market strategist.

Articles of association form a document that specifies the regulations for a company's operations and defines the company's purpose. The document lays out how tasks are to be accomplished within the organization, including the process for appointing directors and the handling of financial records.

Key Takeaways

• Articles of association can be thought of as a user's manual for a company, defining its purpose and outlining the methodology for accomplishing necessary day-to-day tasks.
• The content and terms of the "articles" may vary by jurisdiction, but typically include provisions on the company name, its purpose, the share structure, the company's organization, and provisions concerning shareholder meetings.

In the the U.S. and Canada, articles of association are often referred to as "articles" for short.

Investopedia / Crea Taylor

Articles of association often identify the manner in which a company will issue shares, pay dividends , audit financial records, and provide voting rights . This set of rules can be considered a user's manual for the company because it outlines the methodology for accomplishing the day-to-day tasks that must be completed.

While the content of the articles of association and the exact terms used vary from jurisdiction to jurisdiction, the document is quite similar throughout the world and generally contains provisions on the company name, the company's purpose, the share capital , the company's organization, and provisions regarding shareholder meetings.

Company Name

As a legal entity, the company must have a name that can be found in the articles of association. All jurisdictions will have rules concerning company names. Usually, a suffix such as "Inc." or "Ltd." must be used to show that the entity is a company. Also, some words that could confuse the public, such as "government" or "church," cannot be used or must be used only for specific types of entities. Words that are offensive or heinous are also usually prohibited.

Purpose of the Company

The reason for the creation of the company must also be stated in the articles of association. Some jurisdictions accept very broad purposes—"management"—while others require greater detail—"the operation of a wholesale bakery," for example.

Share Capital

The number and type of shares that comprise a company's capital are listed in the articles of association. There will always be at least one form of common share that makes up a company's capital. In addition, there may be several types of preferred shares . The company may or may not issue the shares , but if they are found in the articles of association, they can be issued if and when the need presents itself.

A company may or may not issue shares, but if they are listed in the articles of association, shares can be issued if and when needed.

Organization of the Company

The legal organization of the company, including its address, the number of directors and officers, and the identity of the founders and original shareholders, are found in this section. Depending on the jurisdiction and type of business, the auditors and legal advisors of the company may also be in this section.

Shareholder Meetings

The provisions for the first general meeting of shareholders and the rules that will govern subsequent annual shareholder meetings —such as notices, resolutions, and votes—are laid out in detail in this section.

A person, or group of people, starting a business will typically refer to a lawyer, accountant , or both for advice when setting up a company.

The company will choose a name and define its purpose. The company is then registered at the state/province or federal level. Note that trademarking a name is a different process.

A company may issue shares to divide up the company if it wishes, but it doesn't need to. The articles will lay out how this can be done. The lawyer or accountant will typically work with the directors of the company, asking them questions to help figure out how they wish to grow and how the company may end up being structured in the future.

Company directors are listed, along with their personal information. A business address is also provided.

Changes can be made to the articles of association with director(s) approval.

Articles of association define a company's purpose as a business, key information such as the name of the business, how the business is organized, and how shares are structured. It is an agreement among members of the organization that defines how business will be conducted.

What Are the Contents of Articles of Association?

The articles of association will include a company's name, its purpose, its share capital, how it is organized, and how shareholders meetings will be conducted. This information will include the names of founders and original shareholders.

Who Creates the Articles of Association?

A company's founders typically will create its articles of association. Because it is an important document addressing the company's management and finances, the creators of the document typically do so in consultation with lawyers, accountants, or both.

Articles of association define what a company is and how it operates. This includes fundamental information such as the company's name, its founders and original shareholders, the purpose driving the company, how shareholders meetings are conducted, and how directors are hired. Company shareholders are bound by the document and use it as a guideline for how to operate the business.

U.S. Small Business Administration. " Register Your Business ."

• Terms of Service
• Editorial Policy
• Privacy Policy
• Your Privacy Choices

• My presentations

Auth with social network:

Download presentation

We think you have liked this presentation. If you wish to download it, please recommend it to your friends in any social system. Share buttons are a little bit lower. Thank you!

Presentation is loading. Please wait.

To view this video please enable JavaScript, and consider upgrading to a web browser that supports HTML5 video

ARTICLES OF ASSOCIATION

Published by Ross Farmer Modified over 7 years ago

Similar presentations

Presentation on theme: "ARTICLES OF ASSOCIATION"— Presentation transcript:

BORROWING POWERS. Capital is necessary for the establishment and development of a business and borrowing is one of the most important source of the capital,

Company act Presented by: Engr.Murtaza zulfiqar.

MEETINGS Shumeet K. Grewal. The word ‘Meetings’ implies the coming together of a certain number of members for transacting the business in agenda, for.

Private and Confidential. Content 1. VIC Constitution 2. Major Parts of Constitution [C1 – C8] 3. Share Capital and Members [C9 – C31] 4. Directors and.

Appointment & Qualifications of Director.  All Private and Public Limited Companies and One Person Companies (OPC)

1 Shareholders (members) 2 Directors 3 Corporate control and minority protection 4 Partner’s Liability.

A NEW CORPORATE FIRM UNDER THE LIMITED LIABILITY PARTERNERSHIP ACT,2008 (Effective from7 th January 2009)

MEETINGS OF SHARE HOLDERS. NEED FOR MEETING TO RATIFY TO EXPRESS THEIR DISAPPROVAL OF, THE DIRECTOR’S PAST CONDUCT. TO CONSIDER THEIR FUTURE PLANS. TO.

Meetings and Resolutions

Memorandum of Association Articles of Association

COURTS, SECP AND REGISTRAR Jurisdiction of Courts Company Benches Procedure of Court Appeals against Court Orders Securities & Exchange Commission of.

BOARD MEETING & COMMITTEE MEETING

Introduction to Company Law

MEMORANDUM OF ASSOCIATION

Corporate Law.

What do you mean by a formation of a Company?

FORMATION OF COMPANIES & THE COMPANY CONSTITUTION CORPORATE LAW.

Articles of Association

1 General meetings including Role of scrutinizers © Savithri Parekh.

About project

© 2024 SlidePlayer.com Inc. All rights reserved.

THE MEMORANDUM AND ARTICLES OF ASSOCIATION

Apr 05, 2019

130 likes | 249 Views

THE MEMORANDUM AND ARTICLES OF ASSOCIATION. Definition of Memo:

Share Presentation

• share certificates
• ultra vires contracts
• company a company
• own government
• certain cases

Presentation Transcript

THE MEMORANDUM AND ARTICLES OF ASSOCIATION Definition of Memo: The purpose of the memorandum is to enable the members of the company, its creditors, and the public to know what its powers are and what is the range of its activities. The memorandum contains rules regarding the capital structure, the liability of the members, the objects of the company, and all other important matters relating to the company. Definition of Articles: The Articles of Association is a document which contains rules, regulations and bye-laws regarding the internal management of the company. Articles must not violate any provision of the memorandum or any provision of the Companies Act. Relationship: Lord Cairns in Riche, described the relationship between the memorandum and the articles in this language: "The memorandum is as it were, the area beyond which the actions of the company cannot go; inside the area, the shareholders may make such regulations for their own government as they think fit". The Articles are subordinate to Memorandum. The Memorandum must be read in conjunction with the Articles. The terms of the Memorandum cannot be modified or controlled by the Articles.

Public Documents: The Memo and Articles are public documents, which may be inspected by anybody at the office of the Registrar of Companies. Differences: • The distinction between the memorandum and the articles of association can be summed up as follows: • The memorandum is determining the company’s constitution and objectives; the articles are rules regarding internal management. • Any rule in the articles contrary to the memorandum is invalid. • Articles can be altered easily, the memorandum can be altered only after the adoption of certain formalities. • Certain clauses of memo cannot be altered without the sanction of the Central Government and of the Court e.g., the object clause and the liability clause. Other clauses can be altered easily e.g., the name clause. Articles can be altered by passing a special resolution. • The memo defines the powers of the company and the relationship between the company and the members and-also non members, Articles define and regulate the relationship between the company and the members. • Acts beyond the powers of memo (ultra vires) are void Such an act cannot be ratified by the members. But acts done by a company beyond the articles can be ratified by the shareholders provided they are within (intra vires) the powers of Memo. • If an act is within the powers given by the memo (irrtra vires the memo) but contrary to some provision of the articles (ultra vires the articles) the members can change the articles and ratify the act.

The form and contents of the Memorandum: The Act lays down that the memorandum of a association of every company shall contain the following particulars : • Name Clause; • Situation Clause; • Objects Clause; • Area of Operation Clause; • Liability Clause; • Capital Clause; • The Association and Subscription Clause Form and contents of the Articles: The Articles of Association contain rules, regulations and bye-laws regarding the internal management of companies. An unlimited company, a company limited by guarantee and a private company limited by shares must file their articles of association at the time of registration of the company.

Form of Articles: The Articles shall: • be printed ; • be divided into paragraphs numbered consecutively ; and • be signed by each subscriber of the memorandum of association, in the presence of at least one witness who shall attest the –signature. Contents of Articles: Articles usually contain provisions in respect of the following matters : • share capital, rights of shareholders, payment of commissions, share certificates; • lien on shares; • calls on shares; • transfer of shares; • transmission of shares; • forfeiture of shares; • conversion of shares into stock; • share warrants; • alteration, of capital; • general meetings and voting rights of members; • appointment and remuneration of directors, board of directors, managers and secretary; • dividends and reserves; • accounts and audit and borrowing powers; • capitalisation of profits; and • winding up.

Alteration of the Memorandum: For the purpose of alteration, the provisions of the memo can be divided into two classes : (i) provisions the inclusion of which is made compulsory by the Act (e.g., the name, objects, place of registered office etc.) (ii) other provisions which the organisers of the -company have thought it desirable to include. Provisions coming under the first category are called "Conditions contained in the Memorandum". The "conditions" can be altered in the manner stated below: 1.Change of name: • A company may change its name by special resolution provided the Company Law Board approves of the change • When the name is validly changed, the Registrar shall enter the new name in the Register of companies and shall issue a fresh Certificate of Incorporation. • Change of name does not affect the rights and obligations of the company and pending suits by or against the company.

2. Change of Object: The object clause of the memo can be changed for the purpose of enabling the company, for example : • to carry on its business more economically or more efficiently; • to attain its main purpose by new or improved means ; • to enlarge or change the local area of its operation; The following procedure must be adopted for changing the object clause : • A special resolution must be passed. • A petition must be filed to the Company Law Board for confirmation of the change. • Notice must be given to all persons whose interests will be affected by the change. • The consent of the creditors of the Company must be obtained or other claims paid off or secured; etc. 3. Change in the location of the registered office from one State to another The procedure to be adopted is the same as in the case of alteration of object. The alteration must be registered with the Registrar of Companies of the 'State in which the registered office of the Company was originally situated and also the Registrar of the 'State to which the office is being transferred.

4. Alteration of the Capital Clause Alteration of the capital clause can be done in the following methods (i) Alteration, including Increase of Capital: (ii) Reduction of Capital (iii) Variation of Shareholders' Rights (iv) Creation of Reserve Capital. Alteration of the Articles Of Association: Although alteration of articles is permitted, there are certain restrictions on the nature and extent of the alterations that can be made, for example: • Articles can be altered by special resolution only. If the articles of the company prescribed a different procedure, e.g., an ordinary resolution, it will not be followed. Confirmation by the Court is not necessary. • No change is permitted which will violate the provisions of the Companies Act. • No change is permitted which is contrary to the conditions contained in the Memorandum of Association of the Company. • The alterations must not 'contain anything illegal. • The liability of the members or any class of members, cannot be increased without their consent. • But any alteration made bona fide, in the interests of the company as a whole, is valid and binding even though the private interests of some members may be affected.

The legal effects of the Memorandum: The Contractual Powers of a Company A Company or a Corporation is a legal person capable of suing and of being sued. But the contractual powers of a company are limited in two ways : (i) natural possibility and (ii) legal possibility. Forms of Contracts and Deeds of a Company • The Doctrine of Ultra Vires • The Memorandum of Association determines the constitution and the powers of 'the Company. • The important rules concerning the legal effects of the memorandum can be summed up as follows, for example: • The terms of the memorandum constitute a binding contract between the Company and the members. • All acts done by the directors or members beyond the powers given in the memo, are ultra vires and not binding on the Company. • The members cannot ratify ultra vires acts, even by an unanimous resolution. • If an act is within the powers given by the memo (intra vires the memo) but contrary to some provision of the articles (ultra vires the articles) members can change the articles and ratify the act. • Directors entering into ultra vires contracts may be liable to the third party for breach of warranty of authority.

Legal effect of the Articles: Binding Contract • The articles constitute a binding contract between the company and its members. • The articles come within the definition of public documents. • All persons dealing with the company are presumed to know the provisions of the articles. So if anything is done contrary to or beyond the provisions of the articles, the company is not bound. Doctrine of indoor management: The Doctrine of indoor management does not apply in certain cases : (a) Void Acts Where the act is void ab initio, the company is not bound, e.g., forgery. (b) Knowledge of irregularity Where the person dealing with the company has notice, actual or constructive, that the prescribed procedure has not been complied with the company is not bound. (c) Lack of authority If an agent of a company makes a contract with a third party and if the act of the agent falls outside the ordinary authority of the agent, the company

• More by User

The Memorandum of Understanding

Yellowstone bison must be allowed to range freely on federal lands to the immediate north and west of the park. ... The bison capture facility within the park must be removed. ...

433 views • 17 slides

The Strecker Memorandum

The Strecker Memorandum. A Review & Commentary Use your arrow keys to navigate by Michael H. Keehn. Intro.

1.67k views • 109 slides

Training Module 3: Articles of Incorporation and Association Bylaws

Training Module 3: Articles of Incorporation and Association Bylaws. Presented by the Southern Early Childhood Association. Topics to be Presented…. Advantages and disadvantages of incorporation Incorporation procedures Articles of Incorporation Association bylaws

729 views • 23 slides

Memorandum of Understanding

Memorandum of Understanding. Col Rafael A Robles, CAP General Counsel. Direct Learning Objectives. What should you take home from this topic? You should be able to: Ascertain when an MOU is appropriate/necessary Understand what alternatives are available to obtain mission authorization

319 views • 15 slides

The Hossbach Memorandum

The Hossbach Memorandum. 5th November, 1937 Hitler’s War Timetable?. At the meeting were Hitler, his adjutant( a kind of personal assistant) called Hossbach, who took notes, and all the heads of the armed forces and foreign policy advisors. . What was the meeting about?.

327 views • 9 slides

Memorandum . Purpose. Flexible form of communication in the organization Function of the letter Effect of a report, briefings, instructions, brief messages Upwards, downwards, side wards Individual to department etc Duplication of memo. Ways of writing. On a blank sheet Memo pads.

374 views • 15 slides

ARTICLES OF ASSOCIATION

ARTICLES OF ASSOCIATION. We assert that we are not powerless over our addiction; that the strength, to cope with and overcome it, lies within us. We assert that it is part of the blueprint of life that each person inherently knows right from wrong:

358 views • 3 slides

Articles of Association

Articles of Association. “ The articles of association is a document regulating the rights of members of the company among themselves and the manner in which the business of the company shall be conducted.” . Forms of Articles of Association.

6k views • 12 slides

ARTICLES OF ASSOCIATION. 1) Articles of Association of a company contain the rules and Regulations relating to the Management of its internal Affairs. 2) It defines the rights, powers and duties of the Management.

603 views • 13 slides

MEMORANDUM. Laws of the Game 2006. Memorandum 2006. 2006 Annual General Meeting International Football Association Board (IFAB) Amendments to the Laws of the Game Instructions and Directives. Memorandum 2006. LAW 4 – The Players’ Equipment.

428 views • 14 slides

Organization of the Office Memorandum

Organization of the Office Memorandum. -IRAC Common Errors/Review -Outlining the Closed Memo -Parts of the Legal Memorandum. IRAC Common Errors. The Issue - should have been fact and law specific:

650 views • 39 slides

Proposed Amendments of the Memorandum & Articles

Proposed Amendments of the Memorandum & Articles. September 28, 2010. Memorandum. Memorandum of Association. Second. “The Registered Office of the Association will be situate situated in the Colony of Hong Kong” (page 5) 更正香港於回歸後的名稱及改正錯字 situate 。

399 views • 20 slides

Memorandum. Ms. Debbie Francisco- Dianco. What is memorandum?. A memorandum or memo is a document or other communication that aids the memory by recording events or observations on a topic, such as may be used in a business office .

1.42k views • 17 slides

Memorandum of Understandings

Memorandum of Understandings. Lt Col Rafael A. Robles, CAP Assistant General Counsel. Direct Learning Objectives. What should you take home from this topic? You should be able to: Ascertain when an MOU is appropriate/necessary

263 views • 15 slides

Memorandum of Understanding. Who What Where When How. And also-Why bother?. Formalize authorities Detail timelines Work out financial considerations Memorialize objectives. An opportunity to Partner. Leverage scarce funds Optimum use of personnel Reduce suspicion

644 views • 12 slides

Memorandum of Understanding. Deakin University (Deakin) and Bangalore University. by Prof. N. Prabhudev Vice-Chancellor Bangalore Unviersity Bangalore. Content. Background Agreement Effective Date and Duration of Agreement Areas of Co-Operation Activities and Projects

254 views • 15 slides

Memorandum Of Incorporation

A Memorandum of Incorporation is one of the key vital that needs to be agreed upon and taken care of for starting up a business in South Africa

43 views • 2 slides

MEMORANDUM OF ASSOCIATION

MEMORANDUM OF ASSOCIATION. Definition Of Memorandum Of Association. According to sec(56) of the companies Act,2013 Memorandum means “Memorandum of association of a company as originally framed or altered from time to time in pursuance of any previous Companies Law or of this act”.

2.22k views • 48 slides

Memorandum of Understanding. Paul E. Reimer 20 June 2008. Purpose. Outlines responsibilities within the collaboration Leadership responsibilities, including safety responsibilities Institutional responsibilities Clearly states who will provide each element of the spectrometer

75 views • 5 slides

• SUGGESTED TOPICS
• The Magazine
• Newsletters
• Managing Yourself
• Managing Teams
• Work-life Balance
• The Big Idea
• Data & Visuals
• Reading Lists
• Case Selections
• HBR Learning
• Topic Feeds
• Account Settings
• Email Preferences

How to Make a “Good” Presentation “Great”

• Guy Kawasaki

Remember: Less is more.

A strong presentation is so much more than information pasted onto a series of slides with fancy backgrounds. Whether you’re pitching an idea, reporting market research, or sharing something else, a great presentation can give you a competitive advantage, and be a powerful tool when aiming to persuade, educate, or inspire others. Here are some unique elements that make a presentation stand out.

• Fonts: Sans Serif fonts such as Helvetica or Arial are preferred for their clean lines, which make them easy to digest at various sizes and distances. Limit the number of font styles to two: one for headings and another for body text, to avoid visual confusion or distractions.
• Colors: Colors can evoke emotions and highlight critical points, but their overuse can lead to a cluttered and confusing presentation. A limited palette of two to three main colors, complemented by a simple background, can help you draw attention to key elements without overwhelming the audience.
• Pictures: Pictures can communicate complex ideas quickly and memorably but choosing the right images is key. Images or pictures should be big (perhaps 20-25% of the page), bold, and have a clear purpose that complements the slide’s text.
• Layout: Don’t overcrowd your slides with too much information. When in doubt, adhere to the principle of simplicity, and aim for a clean and uncluttered layout with plenty of white space around text and images. Think phrases and bullets, not sentences.

As an intern or early career professional, chances are that you’ll be tasked with making or giving a presentation in the near future. Whether you’re pitching an idea, reporting market research, or sharing something else, a great presentation can give you a competitive advantage, and be a powerful tool when aiming to persuade, educate, or inspire others.

• Guy Kawasaki is the chief evangelist at Canva and was the former chief evangelist at Apple. Guy is the author of 16 books including Think Remarkable : 9 Paths to Transform Your Life and Make a Difference.

Partner Center

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

• View all journals
• My Account Login
• Explore content
• About the journal
• Publish with us
• Sign up for alerts
• Open access
• Published: 08 April 2024

Large-scale phenotyping of patients with long COVID post-hospitalization reveals mechanistic subtypes of disease

• Felicity Liew 1   na1 ,
• Claudia Efstathiou   ORCID: orcid.org/0000-0001-6125-8126 1   na1 ,
• Sara Fontanella 1 ,
• Matthew Richardson 2 ,
• Ruth Saunders 2 ,
• Dawid Swieboda 1 ,
• Jasmin K. Sidhu 1 ,
• Stephanie Ascough 1 ,
• Shona C. Moore   ORCID: orcid.org/0000-0001-8610-2806 3 ,
• Noura Mohamed 4 ,
• Jose Nunag   ORCID: orcid.org/0000-0002-4218-0500 5 ,
• Clara King 5 ,
• Olivia C. Leavy 2 , 6 ,
• Omer Elneima 2 ,
• Hamish J. C. McAuley 2 ,
• Aarti Shikotra 7 ,
• Amisha Singapuri   ORCID: orcid.org/0009-0002-4711-7516 2 ,
• Marco Sereno   ORCID: orcid.org/0000-0003-4573-9303 2 ,
• Victoria C. Harris 2 ,
• Linzy Houchen-Wolloff   ORCID: orcid.org/0000-0003-4940-8835 8 ,
• Neil J. Greening   ORCID: orcid.org/0000-0003-0453-7529 2 ,
• Nazir I. Lone   ORCID: orcid.org/0000-0003-2707-2779 9 ,
• Matthew Thorpe 10 ,
• A. A. Roger Thompson   ORCID: orcid.org/0000-0002-0717-4551 11 ,
• Sarah L. Rowland-Jones 11 ,
• Annemarie B. Docherty   ORCID: orcid.org/0000-0001-8277-420X 10 ,
• James D. Chalmers 12 ,
• Ling-Pei Ho   ORCID: orcid.org/0000-0001-8319-301X 13 ,
• Alexander Horsley   ORCID: orcid.org/0000-0003-1828-0058 14 ,
• Betty Raman 15 ,
• Krisnah Poinasamy 16 ,
• Michael Marks 17 , 18 , 19 ,
• Onn Min Kon 1 ,
• Luke S. Howard   ORCID: orcid.org/0000-0003-2822-210X 1 ,
• Daniel G. Wootton 3 ,
• Jennifer K. Quint 1 ,
• Thushan I. de Silva   ORCID: orcid.org/0000-0002-6498-9212 11 ,
• Antonia Ho 20 ,
• Christopher Chiu   ORCID: orcid.org/0000-0003-0914-920X 1 ,
• Ewen M. Harrison   ORCID: orcid.org/0000-0002-5018-3066 10 ,
• William Greenhalf 21 ,
• J. Kenneth Baillie   ORCID: orcid.org/0000-0001-5258-793X 10 , 22 , 23 ,
• Malcolm G. Semple   ORCID: orcid.org/0000-0001-9700-0418 3 , 24 ,
• Lance Turtle 3 , 24 ,
• Rachael A. Evans   ORCID: orcid.org/0000-0002-1667-868X 2 ,
• Louise V. Wain 2 , 6 ,
• Christopher Brightling 2 ,
• Ryan S. Thwaites   ORCID: orcid.org/0000-0003-3052-2793 1   na1 ,
• Peter J. M. Openshaw   ORCID: orcid.org/0000-0002-7220-2555 1   na1 ,
• PHOSP-COVID collaborative group &

ISARIC investigators

Nature Immunology volume  25 ,  pages 607–621 ( 2024 ) Cite this article

25k Accesses

1 Citations

2140 Altmetric

Metrics details

• Inflammasome
• Inflammation
• Innate immunity

One in ten severe acute respiratory syndrome coronavirus 2 infections result in prolonged symptoms termed long coronavirus disease (COVID), yet disease phenotypes and mechanisms are poorly understood 1 . Here we profiled 368 plasma proteins in 657 participants ≥3 months following hospitalization. Of these, 426 had at least one long COVID symptom and 233 had fully recovered. Elevated markers of myeloid inflammation and complement activation were associated with long COVID. IL-1R2, MATN2 and COLEC12 were associated with cardiorespiratory symptoms, fatigue and anxiety/depression; MATN2, CSF3 and C1QA were elevated in gastrointestinal symptoms and C1QA was elevated in cognitive impairment. Additional markers of alterations in nerve tissue repair (SPON-1 and NFASC) were elevated in those with cognitive impairment and SCG3, suggestive of brain–gut axis disturbance, was elevated in gastrointestinal symptoms. Severe acute respiratory syndrome coronavirus 2-specific immunoglobulin G (IgG) was persistently elevated in some individuals with long COVID, but virus was not detected in sputum. Analysis of inflammatory markers in nasal fluids showed no association with symptoms. Our study aimed to understand inflammatory processes that underlie long COVID and was not designed for biomarker discovery. Our findings suggest that specific inflammatory pathways related to tissue damage are implicated in subtypes of long COVID, which might be targeted in future therapeutic trials.

Similar content being viewed by others

Immunopathological signatures in multisystem inflammatory syndrome in children and pediatric COVID-19

Keith Sacco, Riccardo Castagnoli, … Luigi D. Notarangelo

In COVID-19, NLRP3 inflammasome genetic variants are associated with critical disease and these effects are partly mediated by the sickness symptom complex: a nomothetic network approach

Michael Maes, Walton Luiz Del Tedesco Junior, … Andréa Name Colado Simão

Epidemiology, clinical presentation, pathophysiology, and management of long COVID: an update

Sizhen Su, Yimiao Zhao, … Lin Lu

One in ten severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) infections results in post-acute sequelae of coronavirus disease 2019 (PASC) or long coronavirus disease (COVID), which affects 65 million people worldwide 1 . Long COVID (LC) remains common, even after mild acute infection with recent variants 2 , and it is likely LC will continue to cause substantial long-term ill health, requiring targeted management based on an understanding of how disease phenotypes relate to underlying mechanisms. Persistent inflammation has been reported in adults with LC 1 , 3 , but studies have been limited in size, timing of samples or breadth of immune mediators measured, leading to inconsistent or absent associations with symptoms. Markers of oxidative stress, metabolic disturbance, vasculoproliferative processes and IFN-, NF-κB- or monocyte-related inflammation have been suggested 3 , 4 , 5 , 6 .

The PHOSP-COVID study, a multicenter United Kingdom study of patients previously hospitalized with COVID-19, has reported inflammatory profiles in 626 adults with health impairment after COVID-19, identified through clustering. Elevated IL-6 and markers of mucosal inflammation were observed in those with severe impairment compared with individuals with milder impairment 7 . However, LC is a heterogeneous condition that may be a distinct form of health impairment after COVID-19, and it remains unclear whether there are inflammatory changes specific to LC symptom subtypes. Determining whether activated inflammatory pathways underlie all cases of LC or if mechanisms differ according to clinical presentation is essential for developing effective therapies and has been highlighted as a top research priority by patients and clinicians 8 .

In this Letter, in a prospective multicenter study, we measured 368 plasma proteins in 657 adults previously hospitalized for COVID-19 (Fig. 1a and Table 1 ). Individuals in our cohort experienced a range of acute COVID-19 severities based on World Health Organization (WHO) progression scores 9 ; WHO 3–4 (no oxygen support, n  = 133 and median age of 55 years), WHO 5–6 (oxygen support, n  = 353 and median age of 59 years) and WHO 7–9 (critical care, n  = 171 and median age of 57 years). Participants were hospitalized for COVID-19 ≥3 months before sample collection (median 6.1 months, interquartile range (IQR) 5.1–6.8 months and range 3.0–8.3 months) and confirmed clinically ( n  = 36/657) or by PCR ( n  = 621/657). Symptom data indicated 233/657 (35%) felt fully recovered at 6 months (hereafter ‘recovered’) and the remaining 424 (65%) reported symptoms consistent with the WHO definition for LC (symptoms ≥3 months post infection 10 ). Given the diversity of LC presentations, patients were grouped according to symptom type (Fig. 1b ). Groups were defined using symptoms and health deficits that have been commonly reported in the literature 1 ( Methods ). A multivariate penalized logistic regression model (PLR) was used to explore associations of clinical covariates and immune mediators at 6 months between recovered patients ( n  = 233) and each LC group (cardiorespiratory symptoms, cardioresp, n  = 398, Fig. 1c ; fatigue, n  = 384, Fig. 1d ; affective symptoms, anxiety/depression, n  = 202, Fig. 1e ; gastrointestinal symptoms, GI, n  = 132, Fig. 1f ; and cognitive impairment, cognitive, n  = 61, Fig. 1g ). Women ( n  = 239) were more likely to experience CardioResp (odds ratio (OR 1.14), Fatigue (OR 1.22), GI (OR 1.13) and Cognitive (OR 1.03) outcomes (Fig. 1c,d,f,g ). Repeated cross-validation was used to optimize and assess model performance ( Methods and Extended Data Fig. 1 ). Pre-existing conditions, such as chronic lung disease, neurological disease and cardiovascular disease (Supplementary Table 1 ), were associated with all LC groups (Fig. 1c–g ). Age, C-reactive protein (CRP) and acute disease severity were not associated with any LC group (Table 1 ).

a , Distribution of time from COVID-19 hospitalization at sample collection. All samples were cross-sectional. The vertical red line indicates the 3 month cutoff used to define our final cohort and samples collected before 3 months were excluded. b , An UpSet plot describing pooled LC groups. The horizontal colored bars represent the number of patients in each symptom group: cardiorespiratory (Cardio_Resp), fatigue, cognitive, GI and anxiety/depression (Anx_Dep). Vertical black bars represent the number of patients in each symptom combination group. To prevent patient identification, where less than five patients belong to a combination group, this has been represented as ‘<5’. The recovered group ( n  = 233) were used as controls. c – g , Forest plots of Olink protein concentrations (NPX) associated with Cardio_Resp ( n  = 365) ( c ), fatigue (n = 314) ( d ), Anx_Dep ( n  = 202) ( e ), GI ( n  = 124) ( f ) and cognitive ( n  = 60) ( g ). Neuro_Psych, neuropsychiatric. The error bars represent the median accuracy of the model. h , i , Distribution of Olink values (NPX) for IL-1R2 ( h ) and MATN2, neurofascin and sCD58 ( i ) measured between symptomatic and recovered individuals in recovered ( n  = 233), Cardio_Resp ( n  = 365), fatigue ( n  = 314) and Anx_Dep ( n  = 202) groups ( h ) and MATN2 in GI ( n  = 124), neurofascin in cognitive ( n  = 60) and sCD58 in Cardio_Resp and recovered groups ( i ). The box plot center line represents the median, the boundaries represent IQR and the whisker length represents 1.5× IQR. The median values were compared between groups using two-sided Wilcoxon signed-rank test, * P  < 0.05, ** P  < 0.01, *** P  < 0.001 and **** P  < 0.0001.

To study the association of peripheral inflammation with symptoms, we analyzed cross-sectional data collected approximately 6 months after hospitalizations. We measured 368 immune mediators from plasma collected contemporaneously with symptom data. Mediators suggestive of myeloid inflammation were associated with all symptoms (Fig. 1c–h ). Elevated IL-1R2, an IL-1 receptor expressed by monocytes and macrophages modulating inflammation 11 and MATN2, an extracellular matrix protein that modulates tissue inflammation through recruitment of innate immune cells 12 , were associated with cardioresp (IL-1R2 OR 1.14, Fig. 1c,h ), fatigue (IL-1R2 OR 1.45, Fig. 1d,h ), anxiety/depression (IL-1R2 OR 1.34. Fig. 1e,h ) and GI (MATN2 OR 1.08, Fig. 1f ). IL-3RA, an IL-3 receptor, was associated with cardioresp (OR 1.07, Fig. 1c ), fatigue (OR 1.21, Fig. 1d ), anxiety/depression (OR 1.12, Fig. 1e ) and GI (OR 1.06, Fig. 1f ) groups, while CSF3, a cytokine promoting neutrophilic inflammation 13 , was elevated in cardioresp (OR 1.06, Fig. 1c ), fatigue (OR 1.12, Fig. 1d ) and GI (OR 1.08, Fig. 1f ).

Elevated COLEC12, which initiates inflammation in tissues by activating the alternative complement pathway 14 , associated with cardioresp (OR 1.09, Fig. 1c ), fatigue (OR 1.19, Fig. 1d ) and anxiety/depression (OR 1.11, Fig. 1e ), but not with GI (Fig. 1f ) and only weakly with cognitive (OR 1.02, Fig. 1g ). C1QA, a degradation product released by complement activation 15 was associated with GI (OR 1.08, Fig. 1f ) and cognitive (OR 1.03, Fig. 1g ). C1QA, which is known to mediate dementia-related neuroinflammation 16 , had the third strongest association with cognitive (Fig. 1g ). These observations indicated that myeloid inflammation and complement activation were associated with LC.

Increased expression of DPP10 and SCG3 was observed in the GI group compared with recovered (DPP10 OR 1.07 and SCG3 OR 1.08, Fig. 1f ). DPP10 is a membrane protein that modulates tissue inflammation, and increased DPP10 expression is associated with inflammatory bowel disease 17 , 18 , suggesting that GI symptoms may result from enteric inflammation. Elevated SCG3, a multifunctional protein that has been associated with irritable bowel syndrome 19 , suggested that noninflammatory disturbance of the brain–gut axis or dysbiosis, may occur in the GI group. The cognitive group was associated with elevated CTSO (OR 1.04), NFASC (OR 1.03) and SPON-1 (OR 1.02, Fig. 1g,i ). NFASC and SPON-1 regulate neural growth 20 , 21 , while CTSO is a cysteine proteinase supporting tissue turnover 22 . The increased expression of these three proteins as well as C1QA and DPP10 in the cognitive group (Fig. 1g ) suggested neuroinflammation and alterations in nerve tissue repair, possibly resulting in neurodegeneration. Together, our findings indicated that complement activation and myeloid inflammation were common to all LC groups, but subtle differences were observed in the GI and cognitive groups, which may have mechanistic importance. Acutely elevated fibrinogen during hospitalization has been reported to be predictive of LC cognitive deficits 23 . We found elevated fibrinogen in LC relative to recovered (Extended Data Fig. 2a ; P  = 0.0077), although this was not significant when restricted to the cognitive group ( P  = 0.074), supporting our observation of complement pathway activation in LC and in keeping with reports that complement dysregulation and thrombosis drive severe COVID-19 (ref. 24 ).

Elevated sCD58 was associated with lower odds of all LC symptoms and was most pronounced in cardioresp (OR 0.85, Fig. 1c,i ), fatigue (OR 0.80, Fig. 1d ) and anxiety/depression (OR 0.83, Fig. 1e ). IL-2 was negatively associated with the cardioresp (Fig. 1c , OR 0.87), fatigue (Fig. 1d , OR 0.80), anxiety/depression (Fig. 1e , OR 0.84) and cognitive (Fig. 1g , OR 0.96) groups. Both IL-2 and sCD58 have immunoregulatory functions 25 , 26 . Specifically, sCD58 suppresses IL-1- or IL-6-dependent interactions between CD2 + monocytes and CD58 + T or natural killer cells 26 . The association of sCD58 with recovered suggests a central role of dysregulated myeloid inflammation in LC. Elevated markers of tissue repair, IDS and DNER 27 , 28 , were also associated with recovered relative to all LC groups (Fig. 1c–g ). Taken together, our data suggest that suppression of myeloid inflammation and enhanced tissue repair were associated with recovered, supporting the use of immunomodulatory agents in therapeutic trials 29 (Supplementary Table 2 ).

We next sought to validate the experimental and analytical approaches used. Although Olink has been validated against other immunoassay platforms, showing superior sensitivity and specificity 30 , 31 , we confirmed the performance of Olink against chemiluminescent immunoassays within our cohort. We performed chemiluminescent immunoassays on plasma from a subgroup of 58 participants (recovered n  = 13 and LC n  = 45). There were good correlations between results from Olink (normalized protein expression (NPX)) and chemiluminescent immunoassays (pg ml −1 ) for CSF3, IL-1R2, IL-3RA, TNF and TFF2 (Extended Data Fig. 3 ). Most samples did not have concentrations of IL-2 detectable using a mesoscale discovery chemiluminescent assay, limiting this analysis to 14 samples (recovered n  = 4, LC n  = 10, R  = 0.55 and P  = 0.053, Extended Data Fig. 3 ). We next repeated our analysis using alternative definitions of LC. The Centers for Disease Control and Prevention and National Institute for Health and Care Excellence definitions for LC include symptoms occurring 1 month post infection 32 , 33 . Using the 1 month post-infection definition included 62 additional participants to our analysis (recovered n  = 21, 3 females and median age 61 years and LC n  = 41, 15 females and median age 60 years, Extended Data Fig. 2c ) and found that inflammatory associations with each LC group were consistent with our analysis based on the WHO definition (Extended Data Fig. 2d–h ). Finally, to validate the analytical approach (PLR) we examined the distribution of data, prioritizing proteins that were most strongly associated with each LC/recovered group (IL-1R2, MATN2, NFASC and sCD58). Each protein was significantly elevated in the LC group compared with recovered (Fig. 1h,i and Extended Data Fig. 4 ), consistent with the PLR. Alternative regression approaches (unadjusted regression models and partial least squares, PLS) reported results consistent with the original analysis of protein associations and LC outcome in the WHO-defined cohort (Fig. 1c–g , Supplementary Table 3 and Extended Data Figs. 5 and 6 ). The standard errors of PLS estimates were wide (Extended Data Fig. 6 ), consistent with previous demonstrations that PLR is the optimal method to analyze high-dimensional data where variables may have combined effects 34 . As inflammatory proteins are often colinear, working in-tandem to mediate effects, we prioritized PLR results to draw conclusions.

To explore the relationship between inflammatory mediators associated with different LC symptoms, we performed a network analysis of Olink mediators highlighted by PLR within each LC group. COLEC12 and markers of endothelial and mucosal inflammation (MATN2, PCDH1, ROBO1, ISM1, ANGPTL2, TGF-α and TFF2) were highly correlated within the cardioresp, fatigue and anxiety/depression groups (Fig. 2 and Extended Data Fig. 7 ). Elevated PCDH1, an adhesion protein modulating airway inflammation 35 , was highly correlated with other inflammatory proteins associated with the cardioresp group (Fig. 2 ), suggesting that systemic inflammation may arise from the lung in these individuals. This was supported by increased expression of IL-3RA, which regulates innate immune responses in the lung through interactions with circulating IL-3 (ref. 36 ), in fatigue (Figs. 1d and 2 ), which correlated with markers of tissue inflammation, including PCDH1 (Fig. 2 ). MATN2 and ISM1, mucosal proteins that enhance inflammation 37 , 38 , were highly correlated in the GI group (Fig. 2 ), highlighting the role of tissue-specific inflammation in different LC groups. SCG3 correlated less closely with mediators in the GI group (Fig. 2 ), suggesting that the brain–gut axis may contribute separately to some GI symptoms. SPON-1, which regulates neural growth 21 , was the most highly correlated mediator in the cognitive group (Fig. 2 and Extended Data Fig. 7 ), highlighting that processes within nerve tissue may underlie this group. These observations suggested that inflammation might arise from mucosal tissues and that additional mechanisms may contribute to pathophysiology underlying the GI and cognitive groups.

Network analysis of Olink mediators associated with cardioresp ( n  = 365), fatigue ( n  = 314), anxiety/depression ( n  = 202), GI ( n  = 124) and cognitive groups ( n  = 60). Each node corresponds to a protein mediator identified by PLR. The edges (blue lines) were weighted according to the size of Spearman’s rank correlation coefficient between proteins. All edges represent positive and significant correlations ( P  < 0.05) after FDR adjustment.

Women were more likely to experience LC (Table 1 ), as found in previous studies 1 . As estrogen can influence immunological responses 39 , we investigated whether hormonal differences between men and women with LC in our cohort explained this trend. We grouped men and women with LC symptoms into two age groups (those younger than 50 years and those 50 years and older, using age as a proxy for menopause status in women) and compared mediator levels between men and women in each age group, prioritizing those identified by PLR to be higher in LC compared with recovered. As we aimed to understand whether women with LC had stronger inflammatory responses than men with LC, we did not assess differences in men and women in the recovered group. IL-1R2 and MATN2 were significantly higher in women ≥50 years than men ≥50 years in the cardioresp group (Fig. 3a , IL-1R2 and MATN2) and the fatigue group (Fig. 3b ). In the GI group, CSF3 was higher in women ≥50 years compared with men ≥50 years (Fig. 3c ), indicating that the inflammatory markers observed in women were not likely to be estrogen-dependent. Women have been reported to have stronger innate immune responses to infection and to be at greater risk of autoimmunity 39 , possibly explaining why some women in the ≥50 years group had higher inflammatory proteins than men the same group. Proteins associated with the anxiety/depression (IL-1R2 P  = 0.11 and MATN2 P  = 0.61, Extended Data Fig. 8a ) and cognitive groups (CTSO P  = 0.64 and NFASC P  = 0.41, Extended Data Fig. 8b ) were not different between men and women in either age group, consistent with the absent/weak association between sex and these outcomes identified by PLR (Fig. 1e,g ). Though our findings suggested that nonhormonal differences in inflammatory responses may explain why some women are more likely to have LC, they require confirmation in adequately powered studies.

a – c , Olink-measured plasma protein levels (NPX) of IL-1R2 and MATN2 ( a and b ) and CSF3 ( c ) between LC men and LC women divided by age (<50 or ≥50 years) in the cardiorespiratory group (<50 years n  = 8 and ≥50 years n  = 270) ( a ), fatigue group (<50 years n  = 81 and ≥50 years n  = 227) ( b ) and GI group (<50 years n  = 34 and ≥50 years n  = 82) ( c ). the median values were compared between men and women using two-sided Wilcoxon signed-rank test, * P  < 0.05, ** P  < 0.01, *** P  < 0.001 and **** P  < 0.0001. The box plot center line represents the median, the boundaries represent IQR and the whisker length represents 1.5× IQR.

To test whether local respiratory tract inflammation persisted after COVID-19, we compared nasosorption samples from 89 participants (recovered, n  = 31; LC, n  = 33; and healthy SARS-CoV-2 naive controls, n  = 25, Supplementary Tables 4 and 5 ). Several inflammatory markers were elevated in the upper respiratory tract post COVID (including IL-1α, CXCL10, CXCL11, TNF, VEGF and TFF2) when compared with naive controls, but similar between recovered and LC (Fig. 4a ). In the cardioresp group ( n  = 29), inflammatory mediators elevated in plasma (for example, IL-6, APO-2, TGF-α and TFF2) were not elevated in the upper respiratory tract (Extended Data Fig. 9a ) and there was no correlation between plasma and nasal mediator levels (Extended Data Fig. 9b ). This exploratory analysis suggested upper respiratory tract inflammation post COVID was not specifically associated with cardiorespiratory symptoms.

a , Nasal cytokines measured by immunoassay in post-COVID participants ( n  = 64) compared with healthy SARS-CoV-2 naive controls ( n  = 25), and between the the cardioresp group ( n  = 29) and the recovered group ( n  = 31). The red values indicate significantly increased cytokine levels after FDR adjustment ( P  < 0.05) using two-tailed Wilcoxon signed-rank test. b , SARS-CoV-2 N antigen measured in sputum by electrochemiluminescence from recovered ( n  = 17) and pooled LC ( n  = 23) groups, compared with BALF from SARS-CoV-2 naive controls ( n  = 9). The horizontal dashed line indicates the lower limit of detection of the assay. c , Plasma S- and N-specific IgG responses measured by electrochemiluminescence in the LC ( n  = 35) and recovered ( n  = 19) groups. The median values were compared using two-sided Wilcoxon signed-rank tests, NS P  > 0.05, * P  < 0.05, ** P  < 0.01, *** P  < 0.001 and **** P  < 0.0001. The box plot center lines represent the median, the boundaries represent IQR and the whisker length represents 1.5× IQR.

To explore whether SARS-CoV-2 persistence might explain the inflammatory profiles observed in the cardioresp group, we measured SARS-CoV-2 nucleocapsid (N) antigen in sputum from 40 participants (recovered n  = 17 and LC n  = 23) collected approximately 6 months post hospitalization (Supplementary Table 6 ). All samples were compared with prepandemic bronchoalveolar lavage fluid ( n  = 9, Supplementary Table 4 ). Only four samples (recovered n  = 2 and LC n  = 2) had N antigen above the assay’s lower limit of detection, and there was no difference in N antigen concentrations between LC and recovered (Fig. 4b , P  = 0.78). These observations did not exclude viral persistence, which might require tissues samples for detection 40 , 41 . On the basis of the hypothesis that persistent viral antigen might prevent a decline in antibody levels over time, we examined the titers of SARS-CoV-2-specific antibodies in unvaccinated individuals (recovered n  = 19 and LC n  = 35). SARS-CoV-2 N-specific ( P  = 0.023) and spike (S)-specific ( P  = 0.0040) immunoglobulin G (IgG) levels were elevated in LC compared with recovered (Fig. 4c ).

Overall, we identified myeloid inflammation and complement activation in the cardioresp, fatigue, anxiety/depression, cognitive and GI groups 6 months after hospitalization (Extended Data Fig. 10 ). Our findings build on results of smaller studies 5 , 6 , 42 and are consistent with a genome-wide association study that identified an independent association between LC and FOXP4 , which modulates neutrophilic inflammation and immune cell function 43 , 44 . In addition, we identified tissue-specific inflammatory elements, indicating that myeloid disturbance in different tissues may result in distinct symptoms. Multiple mechanisms for LC have been suggested, including autoimmunity, thrombosis, vascular dysfunction, SARS-CoV-2 persistence and latent virus reactivation 1 . All these processes involve myeloid inflammation and complement activation 45 . Complement activation in LC has been suggested in a proteomic study in 97 mostly nonhospitalized COVID-19 cases 42 and a study of 48 LC patients, of which one-third experienced severe acute disease 46 . As components of the complement system are known to have a short half-life 47 , ongoing complement activation suggests active inflammation rather than past tissue damage from acute infection.

Despite the heterogeneity of LC and the likelihood of coexisting or multiple etiologies, our work suggests some common pathways that might be targeted therapeutically and supports the rationale for several drugs currently under trial. Our finding of increased sCD58 levels (associated with suppression of monocyte–lymphocyte interactions 26 ) in the recovered group, strengthens our conclusion that myeloid inflammation is central to the biology of LC and that trials of steroids, IL-1 antagonists, JAK inhibitors, naltrexone and colchicine are justified. Although anticoagulants such as apixaban might prevent thrombosis downstream of complement dysregulation, they can also increase the risk of serious bleeding when given after COVID-19 hospitalization 48 . Thus, clinical trials, already underway, need to carefully assess the risks and benefits of anticoagulants (Supplementary Table 2 ).

Our finding of elevated S- and N-specific IgG in LC could suggest viral persistence, as found in other studies 6 , 42 , 49 . Our network analysis indicated that inflammatory proteins in the cardioresp group interacted strongly with ISM1 and ROBO1, which are expressed during respiratory tract infection and regulate lung inflammation 50 , 51 . Although we were unable to find SARS-CoV-2 antigen in sputum from our LC cases, we did not test for viral persistence in GI tract and lung tissue 40 , 41 or in plasma 52 . Evidence of SARS-CoV-2 persistence would justify trials of antiviral drugs (singly or in combination) in LC. It is also possible that autoimmune processes could result in an innate inflammatory profile in LC. Autoreactive B cells have been identified in LC patients with higher SARS-CoV-2-specific antibody titers in a study of mostly mild acute COVID cases (59% WHO 2–3) 42 , a different population from our study of hospitalized cases.

Our observations of distinct protein profiles in GI and cognitive groups support previous reports on distinct associations between Epstein–Barr virus reactivation and neurological symptoms, or autoantibodies and GI symptoms relative to other forms of LC 49 , 53 . We did not assess autoantibody induction but found evidence of brain–gut axis disturbance (SCG3) in the GI group, which occurs in many autoimmune diseases 54 . We found signatures suggestive of neuroinflammation (C1QA) in the cognitive group, consistent with findings of brain abnormalities on magnetic resonance imaging after COVID-19 hospitalization 55 , as well as findings of microglial activation in mice after COVID-19 (ref. 56 ). Proinflammatory signatures dominated in the cardioresp, fatigue and anxiety/depression groups and were consistent with those seen in non-COVID depression, suggesting shared mechanisms 57 . The association between markers of myeloid inflammation, including IL-3RA, and symptoms was greatest for fatigue. Whilst membrane-bound IL-3RA facilitates IL-3 signaling upstream of myelopoesis 36 its soluble form (measured in plasma) can bind IL-3 and can act as a decoy receptor, preventing monocyte maturation and enhancing immunopathology 58 . Monocytes from individuals with post-COVID fatigue are reported to have abnormal expression profiles (including reduced CXCR2), suggestive of altered maturation and migration 5 , 59 . Lung-specific inflammation was suggested by the association between PCDH1 (an airway epithelial adhesion molecule 35 ) and cardioresp symptoms.

Our observations do not align with all published observations on LC. One proteomic study of 55 LC cases after generally mild (WHO 2–3) acute disease found that TNF and IFN signatures were elevated in LC 3 . Vasculoproliferative processes and metabolic disturbance have been reported in LC 4 , 60 , but these studies used uninfected healthy individuals for comparison and cannot distinguish between LC-specific phenomena and residual post-COVID inflammation. A study of 63 adults (LC, n  = 50 and recovered, n  = 13) reported no association between immune cell activation and LC 3 months after infection 61 , though myeloid inflammation was not directly measured, and 3 months post infection may be too early to detect subtle differences between LC and recovered cases due to residual acute inflammation.

Our study has limitations. We designed the study to identify inflammatory markers identifying pathways underlying LC subgroups rather than diagnostic biomarkers. The ORs we report are small, but associations were consistent across alternative methods of analysis and when using different LC definitions. Small effect sizes can be expected when using PLR, which shrinks correlated mediator coefficients to reflect combined effects and prevent colinear inflation 62 , and could also result from measurement of plasma mediators that may underestimate tissue inflammation. Although our LC cohort is large compared with most other published studies, some of our subgroups are small (only 60 cases were designated cognitive). Though the performance of the cognitive PLR model was adequate, our findings should be validated in larger studies. It should be noted that our cohort of hospitalized cases may not represent all types of LC, especially those occurring after mild infection. We looked for an effect of acute disease severity within our study and did not find it, and are reassured that the inflammatory profiles we observed were consistent with those seen in smaller studies including nonhospitalized cases 42 , 46 . Studies of posthospital LC may be confounded by ‘posthospital syndrome’, which encompasses general and nonspecific effects of hospitalization (particularly intensive care) 63 .

In conclusion, we found markers of myeloid inflammation and complement activation in our large prospective posthospital cohort of patients with LC, in addition to distinct inflammatory patterns in patients with cognitive impairment or gastrointestinal symptoms. These findings show the need to consider subphenotypes in managing patients with LC and support the use of antiviral or immunomodulatory agents in controlled therapeutic trials.

Study design and ethics

After hospitalization for COVID-19, adults who had no comorbidity resulting in a prognosis of less than 6 months were recruited to the PHOSP-COVID study ( n  = 719). Patients hospitalized between February 2020 and January 2021 were recruited. Both sexes were recruited and gender was self-reported (female, n  = 257 and male, n  = 462). Written informed consent was obtained from all patients. Ethical approvals for the PHOSP-COVID study were given by Leeds West Research Ethics Committee (20/YH/0225).

Symptom data and samples were prospectively collected from individuals approximately 6 months (IQR 5.1–6.8 months and range 3.0–8.3 months) post hospitalization (Fig. 1a ), via the PHOSP-COVID multicenter United Kingdom study 64 . Data relating to patient demographics and acute admission were collected via the International Severe Acute Respiratory and Emerging Infection Consortium World Health Organization Clinical Characterisation Protocol United Kingdom (ISARIC4C study; IRAS260007/IRAS126600) (ref. 65 ). Adults hospitalized during the SARS-CoV-2 pandemic were systematically recruited into ISARIC4C. Written informed consent was obtained from all patients. Ethical approval was given by the South Central–Oxford C Research Ethics Committee in England (reference 13:/SC/0149), Scotland A Research Ethics Committee (20/SS/0028) and WHO Ethics Review Committee (RPC571 and RPC572l, 25 April 2013).

Data were collected to account for variables affecting symptom outcome, via hospital records and self-reporting. Acute disease severity was classified according to the WHO clinical progression score: WHO class 3–4: no oxygen therapy; class 5: oxygen therapy; class 6: noninvasive ventilation or high-flow nasal oxygen; and class 7–9: managed in critical care 9 . Clinical data were used to place patients into six categories: ‘recovered’, ‘GI’, ‘cardiorespiratory’, ‘fatigue’, ‘cognitive impairment’ and ‘anxiety/depression’ (Supplementary Table 7 ). Patient-reported symptoms and validated clinical scores were used when feasible, including Medical Research Council (MRC) breathlessness score, dyspnea-12 score, Functional Assessment of Chronic Illness Therapy (FACIT) score, Patient Health Questionnaire (PHQ)-9 and Generalized Anxiety Disorder (GAD)-7. Cognitive impairment was defined as a Montreal Cognitive Assessment score <26. GI symptoms were defined as answering ‘Yes’ to the presence of at least two of the listed symptoms. ‘Recovered’ was defined by self-reporting. Patients were placed in multiple groups if they experienced a combination of symptoms.

Matched nasal fluid and sputum samples were prospectively collected from a subgroup of convalescent patients approximately 6 months after hospitalization via the PHOSP-COVID study. Nasal and bronchoalveolar lavage fluid (BALF) collected from healthy volunteers before the COVID-19 pandemic were used as controls (Supplementary Table 4 ). Written consent was obtained for all individuals and ethical approvals were given by London–Harrow Research Ethics Committee (13/LO/1899) for the collection of nasal samples and the Health Research Authority London–Fulham Research Ethics Committee (IRAS project ID 154109; references 14/LO/1023, 10/H0711/94 and 11/LO/1826) for BALF samples.

Ethylenediaminetetraacetic acid plasma was collected from whole blood taken by venepuncture and frozen at −80 °C as previously described 7 , 66 . Nasal fluid was collected using a NasosorptionTM FX·I device (Hunt Developments), which uses a synthetic absorptive matrix to collect concentrated nasal fluid. Samples were eluted and stored as previously described 67 . Sputum samples were collected via passive expectoration and frozen at −80 °C without the addition of buffers. Sputum samples from convalescent individuals were compared with BALF from healthy SARS-CoV-2-naive controls, collected before the pandemic. BALF samples were used to act as a comparison for lower respiratory tract samples since passively expectorated sputum from healthy SARS-CoV-2-naive individuals was not available. BALF samples were obtained by instillation and recovery of up to 240 ml of normal saline via a fiberoptic bronchoscope. BALF was filtered through 100 µM strainers into sterile 50 ml Falcon tubes, then centrifuged for 10 min at 400  g at 4 °C. The resulting supernatant was transferred into sterile 50 ml Falcon tubes and frozen at −80 °C until use. The full methods for BALF collection and processing have been described previously 68 , 69 .

Immunoassays

To determine inflammatory signatures that associated with symptom outcomes, plasma samples were analyzed on an Olink Explore 384 Inflammation panel 70 . Supplementary Table 8 (Appendix 1 ) lists all the analytes measured. To ensure the validity of results, samples were run in a single batch with the use of negative controls, plate controls in triplicate and repeated measurement of patient samples between plates in duplicate. Samples were randomized between plates according to site and sample collection date. Randomization between plates was blind to LC/recovered outcome. Data were first normalized to an internal extension control that was included in each sample well. Plates were standardized by normalizing to interplate controls, run in triplicate on each plate. Each plate contained a minimum of four patient samples, which were duplicates on another plate; these duplicate pairs allowed any plate to be linked to any other through the duplicates. Data were then intensity normalized across all cohort samples. Finally, Olink results underwent quality control processing and samples or analytes that did not reach quality control standards were excluded. Final normalized relative protein quantities were reported as log 2 NPX values.

To further validate our findings, we performed conventional electrochemiluminescence (ECL) assays and enzyme-linked immunosorbent assay for Olink mediators that were associated with symptom outcome ( Supplementary Methods ). Contemporaneously collected plasma samples were available from 58 individuals. Like most omics platforms, Olink measures relative quantities, so perfect agreement with conventional assays that measure absolute concentrations is not expected.

Sputum samples were thawed before analysis and sputum plugs were extracted with the addition of 0.1% dithiothreitol creating a one in two sample dilution, as previously described 71 . SARS-CoV-2 S and N proteins were measured by ECL S-plex assay at a fixed dilution of one in two (Mesoscale Diagnostics), as per the manufacturers protocol 72 . Control BALF samples were thawed and measured on the same plate, neat. The S-plex assay is highly sensitive in detecting viral antigen in respiratory tract samples 73 .

Nasal cytokines were measured by ECL (mesoscale discovery) and Luminex bead multiplex assays (Biotechne). The full methods and list of analytes are detailed in Supplementary Methods .

Statistics and reproducibility

Clinical data was collected via the PHOSP REDCap database, to which access is available under reasonable request as per the data sharing statement in the manuscript. All analyses were performed within the Outbreak Data Analysis Platform (ODAP). All data and code can be accessed using information in the ‘Data sharing’ and ‘Code sharing’ statements at the end of the manuscript. No statistical method was used to predetermine sample size. Data distribution was assumed to be normal but this was not formally tested. Olink assays and immunoassays were randomized and investigators were blinded to outcomes.

To determine protein signatures that associated with each symptom outcome, a ridge PLR was used. PLR shrinks coefficients to account for combined effects within high-dimensional data, preventing false discovery while managing multicollinearity 34 . Thus, PLR was chosen a priori as the most appropriate model to assess associations between a large number of explanatory variables (that may work together to mediate effects) and symptom outcome 34 , 62 , 70 , 74 . In keeping with our aim to perform an unbiased exploration of inflammatory process, the model alpha was set to zero, facilitating regularization without complete penalization of any mediator. This enabled review of all possible mediators that might associate with LC 62 .

A 50 repeats tenfold nested cross-validation was used to select the optimal lambda for each model and assess its accuracy (Extended Data Fig. 1 ). The performance of the cognitive impairment model was influenced by the imbalance in size of the symptom group ( n  = 60) relative to recovered ( n  = 250). The model was weighted to account for this imbalance resulting in a sensitivity of 0.98, indicating its validity. We have expanded on the model performance and validation approaches in Supplementary Information .

Age, sex, acute disease severity and preexisting comorbidities were included as covariates in the PLR analysis (Supplementary Tables 1 and 3 ). Covariates were selected a priori using features reported to influence the risk of LC and inflammatory responses 1 , 39 , 64 , 75 . Ethnicity was not included since it has been shown not to predict symptom outcome in this cohort 64 . Individuals with missing data were excluded from the regression analysis. Each symptom group was compared with the ‘recovered’ group. The model coefficients of each covariate were converted into ORs for each outcome and visualized in a forest plot, after removing variables associated with regularized OR between 0.98 and 1.02 or in cases where most variables fell outside of this range, using mediators associated with the highest decile of coefficients either side of this range. This enabled exclusion of mediators with effect sizes that were unlikely to have clinical or mechanistic importance since the ridge PLR shrinks and orders coefficients according to their relative importance rather than making estimates with standard error. Thus, confidence intervals cannot be appropriately derived from PLR, and forest plot error bars were calculated using the median accuracy of the model generated by the nested cross-validation. To verify observations made through PLR analysis, we also performed an unadjusted PLR, an unadjusted logistic regression and a PLS analysis. Univariate analyses using Wilcoxon signed-rank test was also performed (Supplementary Table 8 , Appendix 1 ). Analyses were performed in R version 4.2.0 using ‘data.table v1.14.2’, ‘EnvStats v2.7.0’ ‘tidyverse v1.3.2’, ‘lme4 v1.1-32’, ‘caret v6.0-93’, ‘glmnet v4.1-6’, ‘mdatools v0.14.0’, ‘ggpubbr v0.4.0’ and ‘ggplot2 v3.3.6’ packages.

To further investigate the relationship between proteins elevated in each symptom group, we performed a correlation network analysis using Spearman’s rank correlation coefficient and false discovery rate (FDR) thresholding. The mediators visualized in the PLR forest plots, which were associated with cardiorespiratory symptoms, fatigue, anxiety/depression GI symptoms and cognitive impairment were used, respectively. Analyses were performed in R version 4.2.0 using ‘bootnet v1.5.6 ’ and ‘qgraph v1.9.8 ’ packages.

To determine whether differences in protein levels between men and women related to hormonal differences, we divided each symptom group into premenopausal and postmenopausal groups using an age cutoff of 50 years old. Differences between sexes in each group were determined using the Wilcoxon signed-rank test. To understand whether antigen persistence contributed to inflammation in adults with LC, the median viral antigen concentration from sputum/BALF samples and cytokine concentrations from nasal samples were compared using the Wilcoxon signed-rank test. All tests were two-tailed and statistical significance was defined as a P value < 0.05 after adjustment for FDR ( q -value of 0.05). Analyses were performed in R version 4.2.0 using ‘bootnet v1.5.6’ and ‘qgraph v1.9.8’ packages.

Extended Data Fig. 10 was made using Biorender, accessed at www.biorender.com .

Reporting summary

Further information on research design is available in the Nature Portfolio Reporting Summary linked to this article.

Data availability

This is an open access article under the CC BY 4.0 license.

The PHOSP-COVID protocol, consent form, definition and derivation of clinical characteristics and outcomes, training materials, regulatory documents, information about requests for data access, and other relevant study materials are available online at ref. 76 . Access to these materials can be granted by contacting [email protected] and [email protected].

The ISARIC4C protocol, data sharing and publication policy are available at https://isaric4c.net . ISARIC4C’s Independent Data and Material Access Committee welcomes applications for access to data and materials ( https://isaric4c.net ).

The datasets used in the study contain extensive clinical information at an individual level that prevent them from being deposited in an public depository due to data protection policies of the study. Study data can only be accessed via the ODAP, a protected research environment. All data used in this study are available within ODAP and accessible under reasonable request. Data access criteria and information about how to request access is available online at ref. 76 . If criteria are met and a request is made, access can be gained by signing the eDRIS user agreement.

Code availability

Code was written within the ODAP, using R v4.2.0 and publicly available packages (‘data.table v1.14.2’, ‘EnvStats v2.7.0’, ‘tidyverse v1.3.2’, ‘lme4 v1.1-32’, ‘caret v6.0-93’, ‘glmnet v4.1-6’, ‘mdatools v0.14.0’, ‘ggpubbr v0.4.0’, ‘ggplot2 v3.3.6’, ‘bootnet v1.5.6’ and ‘qgraph v1.9.8’ packages). No new algorithms or functions were created and code used in-built functions in listed packages available on CRAN. The code used to generate data and to analyze data is publicly available at https://github.com/isaric4c/wiki/wiki/ISARIC ; https://github.com/SurgicalInformatics/cocin_cc and https://github.com/ClaudiaEfstath/PHOSP_Olink_NatImm .

Davis, H. E., McCorkell, L., Vogel, J. M. & Topol, E. J. Long COVID: major findings, mechanisms and recommendations. Nat. Rev. Microbiol. 21 , 133–146 (2023).

Article   CAS   PubMed   PubMed Central   Google Scholar  

Antonelli, M., Pujol, J. C., Spector, T. D., Ourselin, S. & Steves, C. J. Risk of long COVID associated with delta versus omicron variants of SARS-CoV-2. Lancet 399 , 2263–2264 (2022).

Talla, A. et al. Persistent serum protein signatures define an inflammatory subcategory of long COVID. Nat. Commun. 14 , 3417 (2023).

Captur, G. et al. Plasma proteomic signature predicts who will get persistent symptoms following SARS-CoV-2 infection. EBioMedicine 85 , 104293 (2022).

Scott, N. A. et al. Monocyte migration profiles define disease severity in acute COVID-19 and unique features of long COVID. Eur. Respir. J. https://doi.org/10.1183/13993003.02226-2022 (2023).

Klein, J. et al. Distinguishing features of Long COVID identified through immune profiling. Nature https://doi.org/10.1038/s41586-023-06651-y (2023).

Article   PubMed   PubMed Central   Google Scholar  

Evans, R. A. et al. Clinical characteristics with inflammation profiling of long COVID and association with 1-year recovery following hospitalisation in the UK: a prospective observational study. Lancet Respir. Med . 10 , 761–775 (2022).

Article   CAS   Google Scholar  

Houchen-Wolloff, L. et al. Joint patient and clinician priority setting to identify 10 key research questions regarding the long-term sequelae of COVID-19. Thorax 77 , 717–720 (2022).

Article   PubMed   Google Scholar  

Marshall, J. C. et al. A minimal common outcome measure set for COVID-19 clinical research. Lancet Infect. Dis. 20 , e192–e197 (2020).

Post COVID-19 condition (long COVID). World Health Organization https://www.who.int/europe/news-room/fact-sheets/item/post-covid-19-condition#:~:text=Definition,months%20with%20no%20other%20explanation (2022).

Peters, V. A., Joesting, J. J. & Freund, G. G. IL-1 receptor 2 (IL-1R2) and its role in immune regulation. Brain Behav. Immun. 32 , 1–8 (2013).

Article   CAS   PubMed   Google Scholar  

Luo, Z. et al. Monocytes augment inflammatory responses in human aortic valve interstitial cells via β2-integrin/ICAM-1-mediated signaling. Inflamm. Res. 71 , 681–694 (2022).

Bendall, L. J. & Bradstock, K. F. G-CSF: from granulopoietic stimulant to bone marrow stem cell mobilizing agent. Cytokine Growth Factor Rev. 25 , 355–367 (2014).

Ma, Y. J. et al. Soluble collectin-12 (CL-12) is a pattern recognition molecule initiating complement activation via the alternative pathway. J. Immunol. 195 , 3365–3373 (2015).

Laursen, N. S. et al. Functional and structural characterization of a potent C1q inhibitor targeting the classical pathway of the complement system. Front. Immunol. 11 , 1504 (2020).

Dejanovic, B. et al. Complement C1q-dependent excitatory and inhibitory synapse elimination by astrocytes and microglia in Alzheimer’s disease mouse models. Nat. Aging 2 , 837–850 (2022).

Xue, G., Hua, L., Zhou, N. & Li, J. Characteristics of immune cell infiltration and associated diagnostic biomarkers in ulcerative colitis: results from bioinformatics analysis. Bioengineered 12 , 252–265 (2021).

He, T. et al. Integrative computational approach identifies immune‐relevant biomarkers in ulcerative colitis. FEBS Open Bio. 12 , 500–515 (2022).

Sundin, J. et al. Fecal chromogranins and secretogranins are linked to the fecal and mucosal intestinal bacterial composition of IBS patients and healthy subjects. Sci. Rep. 8 , 16821 (2018).

Kriebel, M., Wuchter, J., Trinks, S. & Volkmer, H. Neurofascin: a switch between neuronal plasticity and stability. Int. J. Biochem. Cell Biol. 44 , 694–697 (2012).

Woo, W.-M. et al. The C. elegans F-spondin family protein SPON-1 maintains cell adhesion in neural and non-neural tissues. Development 135 , 2747–2756 (2008).

Yadati, T., Houben, T., Bitorina, A. & Shiri-Sverdlov, R. The ins and outs of cathepsins: physiological function and role in disease management. Cells 9 , 1679 (2020).

Taquet, M. et al. Acute blood biomarker profiles predict cognitive deficits 6 and 12 months after COVID-19 hospitalization. Nat. Med. https://doi.org/10.1038/s41591-023-02525-y (2023).

Siggins, M. K. et al. Alternative pathway dysregulation in tissues drives sustained complement activation and predicts outcome across the disease course in COVID‐19. Immunology 168 , 473–492 (2023).

Pol, J. G., Caudana, P., Paillet, J., Piaggio, E. & Kroemer, G. Effects of interleukin-2 in immunostimulation and immunosuppression. J. Exp. Med. 217 , e20191247 (2020).

Zhang, Y., Liu, Q., Yang, S. & Liao, Q. CD58 immunobiology at a glance. Front. Immunol. 12 , 705260 (2021).

Demydchuk, M. et al. Insights into Hunter syndrome from the structure of iduronate-2-sulfatase. Nat. Commun. 8 , 15786 (2017).

Wang, Z. et al. DNER promotes epithelial–mesenchymal transition and prevents chemosensitivity through the Wnt/β-catenin pathway in breast cancer. Cell Death Dis. 11 , 642 (2020).

Bonilla, H. et al. Therapeutic trials for long COVID-19: a call to action from the interventions taskforce of the RECOVER initiative. Front. Immunol. 14 , 1129459 (2023).

Wik, L. et al. Proximity extension assay in combination with next-generation sequencing for high-throughput proteome-wide analysis. Mol. Cell. Proteomics 20 , 100168 (2021).

Measuring protein biomarkers with Olink—technical comparisons and orthogonal validation. Olink Proteomics https://www.olink.com/content/uploads/2021/09/olink-technical-comparisons-and-orthogonal-validation-1118-v2.0.pdf (2021).

COVID-19 rapid guideline: managing the long-term effects of COVID-19. National Institute for Health and Care Excellence (NICE), Scottish Intercollegiate Guidelines Network (SIGN) and Royal College of General Practitioners (RCGP) https://www.nice.org.uk/guidance/ng188/resources/covid19-rapid-guideline-managing-the-longterm-effects-of-covid19-pdf-51035515742 (2022).

Long COVID or post-COVID conditions. Centers for Disease Control and Prevention https://www.cdc.gov/coronavirus/2019-ncov/long-term-effects/index.html#:~:text=Long%20COVID%20is%20broadly%20defined,after%20acute%20COVID%2D19%20infection (2023).

Firinguetti, L., Kibria, G. & Araya, R. Study of partial least squares and ridge regression methods. Commun. Stat. Simul. Comput 46 , 6631–6644 (2017).

Article   Google Scholar  

Mortensen, L. J., Kreiner-Moller, E., Hakonarson, H., Bonnelykke, K. & Bisgaard, H. The PCDH1 gene and asthma in early childhood. Eur. Respir. J. 43 , 792–800 (2014).

Tong, Y. et al. The RNFT2/IL-3Rα axis regulates IL-3 signaling and innate immunity. JCI Insight 5 , e133652 (2020).

Wu, Y. et al. Effect of ISM1 on the immune microenvironment and epithelial-mesenchymal transition in colorectal cancer. Front. Cell Dev. Biol. 9 , 681240 (2021).

Luo, G. G. & Ou, J. J. Oncogenic viruses and cancer. Virol. Sin. 30 , 83–84 (2015).

Klein, S. L. & Flanagan, K. L. Sex differences in immune responses. Nat. Rev. Immunol. 16 , 626–638 (2016).

Gaebler, C. et al. Evolution of antibody immunity to SARS-CoV-2. Nature 591 , 639–644 (2021).

Bussani, R. et al. Persistent SARS‐CoV‐2 infection in patients seemingly recovered from COVID‐19. J. Pathol. 259 , 254–263 (2023).

Woodruff, M. C. et al. Chronic inflammation, neutrophil activity, and autoreactivity splits long COVID. Nat. Commun. 14 , 4201 (2023).

Lammi, V. et al. Genome-wide association study of long COVID. Preprint at medRxiv https://doi.org/10.1101/2023.06.29.23292056 (2023).

Ismailova, A. et al. Identification of a forkhead box protein transcriptional network induced in human neutrophils in response to inflammatory stimuli. Front. Immunol. 14 , 1123344 (2023).

Beurskens, F. J., van Schaarenburg, R. A. & Trouw, L. A. C1q, antibodies and anti-C1q autoantibodies. Mol. Immunol. 68 , 6–13 (2015).

Cervia-Hasler, C. et al. Persistent complement dysregulation with signs of thromboinflammation in active long Covid. Science 383 , eadg7942 (2024).

Morgan, B. P. & Harris, C. L. Complement, a target for therapy in inflammatory and degenerative diseases. Nat. Rev. Drug Discov. 14 , 857–877 (2015).

Toshner, M. R. et al. Apixaban following discharge in hospitalised adults with COVID-19: preliminary results from a multicentre, open-label, randomised controlled platform clinical trial. Preprint at medRxiv , https://doi.org/10.1101/2022.12.07.22283175 (2022).

Su, Y. et al. Multiple early factors anticipate post-acute COVID-19 sequelae. Cell 185 , 881–895.e20 (2022).

Branchfield, K. et al. Pulmonary neuroendocrine cells function as airway sensors to control lung immune response. Science 351 , 707–710 (2016).

Rivera-Torruco, G. et al. Isthmin 1 identifies a subset of lung hematopoietic stem cells and it is associated with systemic inflammation. J. Immunol. 202 , 118.18 (2019).

Swank, Z. et al. Persistent circulating severe acute respiratory syndrome coronavirus 2 spike is associated with post-acute coronavirus disease 2019 sequelae. Clin. Infect. Dis. 76 , e487–e490 (2023).

Peluso, M. J. et al. Chronic viral coinfections differentially affect the likelihood of developing long COVID. J. Clin. Invest. 133 , e163669 (2023).

Bellocchi, C. et al. The interplay between autonomic nervous system and inflammation across systemic autoimmune diseases. Int. J. Mol. Sci. 23 , 2449 (2022).

Raman, B. et al. Multiorgan MRI findings after hospitalisation with COVID-19 in the UK (C-MORE): a prospective, multicentre, observational cohort study. Lancet Respir. Med 11 , 1003–1019 (2023).

Fernández-Castañeda, A. et al. Mild respiratory COVID can cause multi-lineage neural cell and myelin dysregulation. Cell 185 , 2452–2468.e16 (2022).

Dantzer, R., O’Connor, J. C., Freund, G. G., Johnson, R. W. & Kelley, K. W. From inflammation to sickness and depression: when the immune system subjugates the brain. Nat. Rev. Neurosci. 9 , 46–56 (2008).

Broughton, S. E. et al. Dual mechanism of interleukin-3 receptor blockade by an anti-cancer antibody. Cell Rep. 8 , 410–419 (2014).

Ley, K., Miller, Y. I. & Hedrick, C. C. Monocyte and macrophage dynamics during atherogenesis. Arterioscler. Thromb. Vasc. Biol. 31 , 1506–1516 (2011).

Iosef, C. et al. Plasma proteome of long-COVID patients indicates HIF-mediated vasculo-proliferative disease with impact on brain and heart function. J. Transl. Med. 21 , 377 (2023).

Santopaolo, M. et al. Prolonged T-cell activation and long COVID symptoms independently associate with severe COVID-19 at 3 months. eLife 12 , e85009 (2023).

Friedman, J., Hastie, T. & Tibshirani, R. Regularization paths for generalized linear models via coordinate descent. J. Stat. Softw. 33 , 1–22 (2010).

Voiriot, G. et al. Chronic critical illness and post-intensive care syndrome: from pathophysiology to clinical challenges. Ann. Intensive Care 12 , 58 (2022).

Evans, R. A. et al. Physical, cognitive, and mental health impacts of COVID-19 after hospitalisation (PHOSP-COVID): a UK multicentre, prospective cohort study. Lancet Respir. Med 9 , 1275–1287 (2021).

Docherty, A. B. et al. Features of 20,133 UK patients in hospital with covid-19 using the ISARIC WHO clinical characterisation protocol: prospective observational cohort study. BMJ https://doi.org/10.1136/bmj.m1985 (2020).

Elneima, O. et al. Cohort profile: post-hospitalisation COVID-19 study (PHOSP-COVID). Preprint at medRxiv https://doi.org/10.1101/2023.05.08.23289442 (2023).

Liew, F. et al. SARS-CoV-2-specific nasal IgA wanes 9 months after hospitalisation with COVID-19 and is not induced by subsequent vaccination. EBioMedicine 87 , 104402 (2023).

Ascough, S. et al. Divergent age-related humoral correlates of protection against respiratory syncytial virus infection in older and young adults: a pilot, controlled, human infection challenge model. Lancet Healthy Longev. 3 , e405–e416 (2022).

Guvenel, A. et al. Epitope-specific airway-resident CD4+ T cell dynamics during experimental human RSV infection. J. Clin. Invest. 130 , 523–538 (2019).

Article   PubMed Central   Google Scholar  

Greenwood, C. J. et al. A comparison of penalised regression methods for informing the selection of predictive markers. PLoS ONE 15 , e0242730 (2020).

Higham, A. et al. Leukotriene B4 levels in sputum from asthma patients. ERJ Open Res. 2 , 00088–02015 (2016).

SARS-CoV-2 spike kit. MSD https://www.mesoscale.com/~/media/files/product%20inserts/s-plex%20sars-cov-2%20spike%20kit%20product%20insert.pdf (2023).

Ren, A. et al. Ultrasensitive assay for saliva-based SARS-CoV-2 antigen detection. Clin. Chem. Lab. Med. 60 , 771–777 (2022).

Breheny, P. & Huang, J. Penalized methods for bi-level variable selection. Stat. Interface 2 , 369–380 (2009).

Thwaites, R. S. et al. Inflammatory profiles across the spectrum of disease reveal a distinct role for GM-CSF in severe COVID-19. Sci. Immunol. 6 , eabg9873 (2021).

Resources. PHOSP-COVID https://phosp.org/resource/ (2022).

Download references

Acknowledgements

This research used data assets made available by ODAP as part of the Data and Connectivity National Core Study, led by Health Data Research UK in partnership with the Office for National Statistics and funded by UK Research and Innovation (grant ref. MC_PC_20058). This work is supported by the following grants: the PHOSP-COVD study is jointly funded by UK Research and Innovation and National Institute of Health and Care Research (NIHR; grant references MR/V027859/1 and COV0319). ISARIC4C is supported by grants from the National Institute for Health and Care Research (award CO-CIN-01) and the MRC (grant MC_PC_19059) Liverpool Experimental Cancer Medicine Centre provided infrastructure support for this research (grant reference C18616/A25153). Other grants that have supported this work include the UK Coronavirus Immunology Consortium (funder reference 1257927), the Imperial Biomedical Research Centre (NIHR Imperial BRC, grant IS-BRC-1215-20013), the Health Protection Research Unit in Respiratory Infections at Imperial College London and NIHR Health Protection Research Unit in Emerging and Zoonotic Infections at University of Liverpool, both in partnership with Public Health England, (NIHR award 200907), Wellcome Trust and Department for International Development (215091/Z/18/Z), Health Data Research UK (grant code 2021.0155), MRC (grant code MC_UU_12014/12) and NIHR Clinical Research Network for providing infrastructure support for this research. We also acknowledge the support of the MRC EMINENT Network (MR/R502121/1), which is cofunded by GSK, the Comprehensive Local Research Networks, the MRC HIC-Vac network (MR/R005982/1) and the RSV Consortium in Europe Horizon 2020 Framework Grant 116019. F.L. is supported by an MRC clinical training fellowship (award MR/W000970/1). C.E. is funded by NIHR (grant P91258-4). L.-P.H. is supported by Oxford NIHR Biomedical Research Centre. A.A.R.T. is supported by a British Heart Foundation (BHF) Intermediate Clinical Fellowship (FS/18/13/33281). S.L.R.-J. receives support from UK Research and Innovation (UKRI), Global Challenges Research Fund (GCRF), Rosetrees Trust, British HIV association (BHIVA), European & Developing Countries Clinical Trials Partnership (EDCTP) and Globvac. J.D.C. has grants from AstraZeneca, Boehringer Ingelheim, GSK, Gilead Sciences, Grifols, Novartis and Insmed. R.A.E. holds a NIHR Clinician Scientist Fellowship (CS-2016-16-020). A. Horsley is currently supported by UK Research and Innovation, NIHR and NIHR Manchester BRC. B.R. receives support from BHF Oxford Centre of Research Excellence, NIHR Oxford BRC and MRC. D.G.W. is supported by an NIHR Advanced Fellowship. A. Ho has received support from MRC and for the Coronavirus Immunology Consortium (MR/V028448/1). L.T. is supported by the US Food and Drug Administration Medical Countermeasures Initiative contract 75F40120C00085 and the National Institute for Health Research Health Protection Research Unit in Emerging and Zoonotic Infections (NIHR200907) at the University of Liverpool in partnership with UK Health Security Agency (UK-HSA), in collaboration with Liverpool School of Tropical Medicine and the University of Oxford. L.V.W. has received support from UKRI, GSK/Asthma and Lung UK and NIHR for this study. M.G.S. has received support from NIHR UK, MRC UK and Health Protection Research Unit in Emerging and Zoonotic Infections, University of Liverpool. J.K.B. is supported by the Wellcome Trust (223164/Z/21/Z) and UKRI (MC_PC_20004, MC_PC_19025, MC_PC_1905, MRNO2995X/1 and MC_PC_20029). The funders were not involved in the study design, interpretation of data or writing of this manuscript. The views expressed are those of the authors and not necessarily those of the Department of Health and Social Care (DHSC), the Department for International Development (DID), NIHR, MRC, the Wellcome Trust, UK-HSA, the National Health Service or the Department of Health. P.J.M.O. is supported by a NIHR Senior Investigator Award (award 201385). We thank all the participants and their families. We thank the many research administrators, health-care and social-care professionals who contributed to setting up and delivering the PHOSP-COVID study at all of the 65 NHS trusts/health boards and 25 research institutions across the United Kingdom, as well as those who contributed to setting up and delivering the ISARIC4C study at 305 NHS trusts/health boards. We also thank all the supporting staff at the NIHR Clinical Research Network, Health Research Authority, Research Ethics Committee, Department of Health and Social Care, Public Health Scotland and Public Health England. We thank K. Holmes at the NIHR Office for Clinical Research Infrastructure for her support in coordinating the charities group. The PHOSP-COVID industry framework was formed to provide advice and support in commercial discussions, and we thank the Association of the British Pharmaceutical Industry as well the NIHR Office for Clinical Research Infrastructure for coordinating this. We are very grateful to all the charities that have provided insight to the study: Action Pulmonary Fibrosis, Alzheimer’s Research UK, Asthma and Lung UK, British Heart Foundation, Diabetes UK, Cystic Fibrosis Trust, Kidney Research UK, MQ Mental Health, Muscular Dystrophy UK, Stroke Association Blood Cancer UK, McPin Foundations and Versus Arthritis. We thank the NIHR Leicester Biomedical Research Centre patient and public involvement group and Long Covid Support. We also thank G. Khandaker and D. C. Newcomb who provided valuable feedback on this work. Extended Data Fig. 10 was created using Biorender.

Author information

These authors contributed equally: Felicity Liew, Claudia Efstathiou, Ryan S. Thwaites, Peter J. M. Openshaw.

Authors and Affiliations

National Heart and Lung Institute, Imperial College London, London, UK

Felicity Liew, Claudia Efstathiou, Sara Fontanella, Dawid Swieboda, Jasmin K. Sidhu, Stephanie Ascough, Onn Min Kon, Luke S. Howard, Jennifer K. Quint, Christopher Chiu, Ryan S. Thwaites, Peter J. M. Openshaw, Jake Dunning & Peter J. M. Openshaw

Institute for Lung Health, Leicester NIHR Biomedical Research Centre, University of Leicester, Leicester, UK

Matthew Richardson, Ruth Saunders, Olivia C. Leavy, Omer Elneima, Hamish J. C. McAuley, Amisha Singapuri, Marco Sereno, Victoria C. Harris, Neil J. Greening, Rachael A. Evans, Louise V. Wain, Christopher Brightling & Ananga Singapuri

NIHR Health Protection Research Unit in Emerging and Zoonotic Infections, Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool, UK

Shona C. Moore, Daniel G. Wootton, Malcolm G. Semple, Lance Turtle, William A. Paxton & Georgios Pollakis

The Imperial Clinical Respiratory Research Unit, Imperial College NHS Trust, London, UK

Noura Mohamed

Cardiovascular Research Team, Imperial College Healthcare NHS Trust, London, UK

Jose Nunag & Clara King

Department of Population Health Sciences, University of Leicester, Leicester, UK

Olivia C. Leavy, Louise V. Wain & Beatriz Guillen-Guio

NIHR Leicester Biomedical Research Centre, University of Leicester, Leicester, UK

Aarti Shikotra

Centre for Exercise and Rehabilitation Science, NIHR Leicester Biomedical Research Centre-Respiratory, University of Leicester, Leicester, UK

Linzy Houchen-Wolloff

Usher Institute, University of Edinburgh, Edinburgh, UK

Nazir I. Lone, Luke Daines, Annemarie B. Docherty, Nazir I. Lone, Matthew Thorpe, Annemarie B. Docherty, Thomas M. Drake, Cameron J. Fairfield, Ewen M. Harrison, Stephen R. Knight, Kenneth A. Mclean, Derek Murphy, Lisa Norman, Riinu Pius & Catherine A. Shaw

Centre for Medical Informatics, The Usher Institute, University of Edinburgh, Edinburgh, UK

Matthew Thorpe, Annemarie B. Docherty, Ewen M. Harrison, J. Kenneth Baillie, Sarah L. Rowland-Jones, A. A. Roger Thompson & Thushan de Silva

Department of Infection, Immunity and Cardiovascular Disease, University of Sheffield, Sheffield, UK

A. A. Roger Thompson, Sarah L. Rowland-Jones, Thushan I. de Silva & James D. Chalmers

University of Dundee, Ninewells Hospital and Medical School, Dundee, UK

James D. Chalmers & Ling-Pei Ho

MRC Human Immunology Unit, University of Oxford, Oxford, UK

Ling-Pei Ho & Alexander Horsley

Division of Infection, Immunity and Respiratory Medicine, Faculty of Biology, Medicine and Health, University of Manchester, Manchester, UK

Alexander Horsley & Betty Raman

Radcliffe Department of Medicine, University of Oxford, Oxford, UK

Betty Raman & Krisnah Poinasamy

Asthma + Lung UK, London, UK

Krisnah Poinasamy & Michael Marks

Department of Clinical Research, London School of Hygiene and Tropical Medicine, London, UK

Michael Marks

Hospital for Tropical Diseases, University College London Hospital, London, UK

Division of Infection and Immunity, University College London, London, UK

Michael Marks & Mahdad Noursadeghi

MRC Centre for Virus Research, School of Infection and Immunity, University of Glasgow, Glasgow, UK

Antonia Ho & William Greenhalf

Institute of Systems, Molecular and Integrative Biology, University of Liverpool, Liverpool, UK

William Greenhalf & J. Kenneth Baillie

The Roslin Institute, University of Edinburgh, Edinburgh, UK

J. Kenneth Baillie, J. Kenneth Baillie, Sara Clohisey, Fiona Griffiths, Ross Hendry, Andrew Law & Wilna Oosthuyzen

Pandemic Science Hub, University of Edinburgh, Edinburgh, UK

J. Kenneth Baillie

The Pandemic Institute, University of Liverpool, Liverpool, UK

Malcolm G. Semple & Lance Turtle

University of Manchester, Manchester, UK

Kathryn Abel, Perdita Barran, H. Chinoy, Bill Deakin, M. Harvie, C. A. Miller, Stefan Stanel & Drupad Trivedi

Intensive Care Unit, Royal Infirmary of Edinburgh, Edinburgh, UK

Kathryn Abel & J. Kenneth Baillie

North Bristol NHS Trust and University of Bristol, Bristol, UK

H. Adamali, David Arnold, Shaney Barratt, A. Dipper, Sarah Dunn, Nick Maskell, Anna Morley, Leigh Morrison, Louise Stadon, Samuel Waterson & H. Welch

University of Edinburgh, Manchester, UK

Davies Adeloye, D. E. Newby, Riinu Pius, Igor Rudan, Manu Shankar-Hari, Catherine Sudlow, Sarah Walmsley & Bang Zheng

King’s College Hospital NHS Foundation Trust and King’s College London, London, UK

Oluwaseun Adeyemi, Rita Adrego, Hosanna Assefa-Kebede, Jonathon Breeze, S. Byrne, Pearl Dulawan, Amy Hoare, Caroline Jolley, Abigail Knighton, M. Malim, Sheetal Patale, Ida Peralta, Natassia Powell, Albert Ramos, K. Shevket, Fabio Speranza & Amelie Te

Guy’s and St Thomas’ NHS Foundation Trust, London, UK

Laura Aguilar Jimenez, Gill Arbane, Sarah Betts, Karen Bisnauthsing, A. Dewar, Nicholas Hart, G. Kaltsakas, Helen Kerslake, Murphy Magtoto, Philip Marino, L. M. Martinez, Marlies Ostermann, Jennifer Rossdale & Teresa Solano

Royal Free London NHS Foundation Trust, London, UK

Shanaz Ahmad, Simon Brill, John Hurst, Hannah Jarvis, C. Laing, Lai Lim, S. Mandal, Darwin Matila, Olaoluwa Olaosebikan & Claire Singh

University Hospital Birmingham NHS Foundation Trust and University of Birmingham, Birmingham, UK

N. Ahmad Haider, Catherine Atkin, Rhiannon Baggott, Michelle Bates, A. Botkai, Anna Casey, B. Cooper, Joanne Dasgin, Camilla Dawson, Katharine Draxlbauer, N. Gautam, J. Hazeldine, T. Hiwot, Sophie Holden, Karen Isaacs, T. Jackson, Vicky Kamwa, D. Lewis, Janet Lord, S. Madathil, C. McGee, K. Mcgee, Aoife Neal, Alex Newton-Cox, Joseph Nyaboko, Dhruv Parekh, Z. Peterkin, H. Qureshi, Liz Ratcliffe, Elizabeth Sapey, J. Short, Tracy Soulsby, J. Stockley, Zehra Suleiman, Tamika Thompson, Maximina Ventura, Sinead Walder, Carly Welch, Daisy Wilson, S. Yasmin & Kay Por Yip

Stroke Association, London, UK

Rubina Ahmed & Richard Francis

University College London Hospital and University College London, London, UK

Nyarko Ahwireng, Dongchun Bang, Donna Basire, Jeremy Brown, Rachel Chambers, A. Checkley, R. Evans, M. Heightman, T. Hillman, Joseph Jacob, Roman Jastrub, M. Lipman, S. Logan, D. Lomas, Marta Merida Morillas, Hannah Plant, Joanna Porter, K. Roy & E. Wall

Oxford University Hospitals NHS Foundation Trust and University of Oxford, Oxford, UK

Mark Ainsworth, Asma Alamoudi, Angela Bloss, Penny Carter, M. Cassar, Jin Chen, Florence Conneh, T. Dong, Ranuromanana Evans, V. Ferreira, Emily Fraser, John Geddes, F. Gleeson, Paul Harrison, May Havinden-Williams, P. Jezzard, Ivan Koychev, Prathiba Kurupati, H. McShane, Clare Megson, Stefan Neubauer, Debby Nicoll, C. Nikolaidou, G. Ogg, Edmund Pacpaco, M. Pavlides, Yanchun Peng, Nayia Petousi, John Pimm, Najib Rahman, M. J. Rowland, Kathryn Saunders, Michael Sharpe, Nick Talbot, E. M. Tunnicliffe & C. Xie

St George’s University Hospitals NHS Foundation Trust, London, UK

Mariam Ali, Raminder Aul, A. Dunleavy, D. Forton, Mark Mencias, N. Msimanga, T. Samakomva, Sulman Siddique, Vera Tavoukjian & J. Teixeira

University Hospitals of Leicester NHS Trust and University of Leicester, Leicester, UK

M. Aljaroof, Natalie Armstrong, H. Arnold, Hnin Aung, Majda Bakali, M. Bakau, E. Baldry, Molly Baldwin, Charlotte Bourne, Michelle Bourne, Nigel Brunskill, P. Cairns, Liesel Carr, Amanda Charalambou, C. Christie, Melanie Davies, Enya Daynes, Sarah Diver, Rachael Dowling, Sarah Edwards, C. Edwardson, H. Evans, J. Finch, Sarah Glover, Nicola Goodman, Bibek Gooptu, Kate Hadley, Pranab Haldar, Beverley Hargadon, W. Ibrahim, L. Ingram, Kamlesh Khunti, A. Lea, D. Lee, Gerry McCann, P. McCourt, Teresa Mcnally, George Mills, Will Monteiro, Manish Pareek, S. Parker, Anne Prickett, I. N. Qureshi, A. Rowland, Richard Russell, Salman Siddiqui, Sally Singh, J. Skeemer, M. Soares, E. Stringer, T. Thornton, Martin Tobin, T. J. C. Ward, F. Woodhead, Tom Yates & A. J. Yousuf

University of Exeter, Exeter, UK

Louise Allan, Clive Ballard & Andrew McGovern

University of Leicester, Leicester, UK

Richard Allen, Michelle Bingham, Terry Brugha, Selina Finney, Rob Free, Don Jones, Claire Lawson, Daniel Lozano-Rojas, Gardiner Lucy, Alistair Moss, Elizabeta Mukaetova-Ladinska, Petr Novotny, Kimon Ntotsis, Charlotte Overton, John Pearl, Tatiana Plekhanova, M. Richardson, Nilesh Samani, Jack Sargant, Ruth Saunders, M. Sharma, Mike Steiner, Chris Taylor, Sarah Terry, C. Tong, E. Turner, J. Wormleighton & Bang Zhao

Liverpool University Hospitals NHS Foundation Trust and University of Liverpool, Liverpool, UK

Lisa Allerton, Ann Marie Allt, M. Beadsworth, Anthony Berridge, Jo Brown, Shirley Cooper, Andy Cross, Sylviane Defres, S. L. Dobson, Joanne Earley, N. French, Kera Hainey, Hayley Hardwick, Jenny Hawkes, Victoria Highett, Sabina Kaprowska, Angela Key, Lara Lavelle-Langham, N. Lewis-Burke, Gladys Madzamba, Flora Malein, Sophie Marsh, Chloe Mears, Lucy Melling, Matthew Noonan, L. Poll, James Pratt, Emma Richardson, Anna Rowe, Victoria Shaw, K. A. Tripp, Lilian Wajero, S. A. Williams-Howard, Dan Wootton & J. Wyles

Sherwood Forest Hospitals NHS Foundation Trust, Nottingham, UK

Lynne Allsop, Kaytie Bennett, Phil Buckley, Margaret Flynn, Mandy Gill, Camelia Goodwin, M. Greatorex, Heidi Gregory, Cheryl Heeley, Leah Holloway, Megan Holmes, John Hutchinson, Jill Kirk, Wayne Lovegrove, Terri Ann Sewell, Sarah Shelton, D. Sissons, Katie Slack, Susan Smith, D. Sowter, Sarah Turner, V. Whitworth & Inez Wynter

Nottingham University Hospitals NHS Trust and University of Nottingham, London, UK

Paula Almeida, Akram Hosseini, Robert Needham & Karen Shaw

Manchester University NHS Foundation Trust and University of Manchester, London, UK

Bashar Al-Sheklly, Cristina Avram, John Blaikely, M. Buch, N. Choudhury, David Faluyi, T. Felton, T. Gorsuch, Neil Hanley, Tracy Hussell, Zunaira Kausar, Natasha Odell, Rebecca Osbourne, Karen Piper Hanley, K. Radhakrishnan & Sue Stockdale

Imperial College London, London, UK

Danny Altmann, Anew Frankel, Luke S. Howard, Desmond Johnston, Liz Lightstone, Anne Lingford-Hughes, William Man, Steve McAdoo, Jane Mitchell, Philip Molyneaux, Christos Nicolaou, D. P. O’Regan, L. Price, Jennifer K. Quint, David Smith, Jonathon Valabhji, Simon Walsh, Martin Wilkins & Michelle Willicombe

Hampshire Hospitals NHS Foundation Trust, Basingstoke, UK

Maria Alvarez Corral, Ava Maria Arias, Emily Bevan, Denise Griffin, Jane Martin, J. Owen, Sheila Payne, A. Prabhu, Annabel Reed, Will Storrar, Nick Williams & Caroline Wrey Brown

British Heart Foundation, Birmingham, UK

Shannon Amoils

NHS Greater Glasgow and Clyde Health Board and University of Glasgow, Glasgow, UK

David Anderson, Neil Basu, Hannah Bayes, Colin Berry, Ammani Brown, Andrew Dougherty, K. Fallon, L. Gilmour, D. Grieve, K. Mangion, I. B. McInnes, A. Morrow, Kathryn Scott & R. Sykes

University of Oxford, Oxford, UK

Charalambos Antoniades, A. Bates, M. Beggs, Kamaldeep Bhui, Katie Breeze, K. M. Channon, David Clark, X. Fu, Masud Husain, Lucy Kingham, Paul Klenerman, Hanan Lamlum, X. Li, E. Lukaschuk, Celeste McCracken, K. McGlynn, R. Menke, K. Motohashi, T. E. Nichols, Godwin Ogbole, S. Piechnik, I. Propescu, J. Propescu, A. A. Samat, Z. B. Sanders, Louise Sigfrid & M. Webster

Belfast Health and Social Care Trust and Queen’s University Belfast, Belfast, UK

Cherie Armour, Vanessa Brown, John Busby, Bronwen Connolly, Thelma Craig, Stephen Drain, Liam Heaney, Bernie King, Nick Magee, E. Major, Danny McAulay, Lorcan McGarvey, Jade McGinness, Tunde Peto & Roisin Stone

Airedale NHS Foundation Trust, Keighley, UK

Lisa Armstrong, Brigid Hairsine, Helen Henson, Claire Kurasz, Alison Shaw & Liz Shenton

Wrightington Wigan and Leigh NHS Trust, Wigan, UK

A. Ashish, Josh Cooper & Emma Robinson

Leeds Teaching Hospitals and University of Leeds, Leeds, UK

Andrew Ashworth, Paul Beirne, Jude Clarke, C. Coupland, Matthhew Dalton, Clair Favager, Jodie Glossop, John Greenwood, Lucy Hall, Tim Hardy, Amy Humphries, Jennifer Murira, Dan Peckham, S. Plein, Jade Rangeley, Gwen Saalmink, Ai Lyn Tan, Elaine Wade, Beverley Whittam, Nicola Window & Janet Woods

University of Liverpool, Liverpool, UK

M. Ashworth, D. Cuthbertson, G. Kemp, Anne McArdle, Benedict Michael, Will Reynolds, Lisa Spencer, Ben Vinson, Katie A. Ahmed, Jane A. Armstrong, Milton Ashworth, Innocent G. Asiimwe, Siddharth Bakshi, Samantha L. Barlow, Laura Booth, Benjamin Brennan, Katie Bullock, Nicola Carlucci, Emily Cass, Benjamin W. A. Catterall, Jordan J. Clark, Emily A. Clarke, Sarah Cole, Louise Cooper, Helen Cox, Christopher Davis, Oslem Dincarslan, Alejandra Doce Carracedo, Chris Dunn, Philip Dyer, Angela Elliott, Anthony Evans, Lorna Finch, Lewis W. S. Fisher, Lisa Flaherty, Terry Foster, Isabel Garcia-Dorival, Philip Gunning, Catherine Hartley, Karl Holden, Anthony Holmes, Rebecca L. Jensen, Christopher B. Jones, Trevor R. Jones, Shadia Khandaker, Katharine King, Robyn T. Kiy, Chrysa Koukorava, Annette Lake, Suzannah Lant, Diane Latawiec, Lara Lavelle-Langham, Daniella Lefteri, Lauren Lett, Lucia A. Livoti, Maria Mancini, Hannah Massey, Nicole Maziere, Sarah McDonald, Laurence McEvoy, John McLauchlan, Soeren Metelmann, Nahida S. Miah, Joanna Middleton, Joyce Mitchell, Ellen G. Murphy, Rebekah Penrice-Randal, Jack Pilgrim, Tessa Prince, P. Matthew Ridley, Debby Sales, Rebecca K. Shears, Benjamin Small, Krishanthi S. Subramaniam, Agnieska Szemiel, Aislynn Taggart, Jolanta Tanianis-Hughes, Jordan Thomas, Erwan Trochu, Libby van Tonder, Eve Wilcock & J. Eunice Zhang

University College London, London, UK

Shahab Aslani, Amita Banerjee, R. Batterham, Gabrielle Baxter, Robert Bell, Anthony David, Emma Denneny, Alun Hughes, W. Lilaonitkul, P. Mehta, Ashkan Pakzad, Bojidar Rangelov, B. Williams, James Willoughby & Moucheng Xu

Hull University Teaching Hospitals NHS Trust and University of Hull, Hull, UK

Paul Atkin, K. Brindle, Michael Crooks, Katie Drury, Nicholas Easom, Rachel Flockton, L. Holdsworth, A. Richards, D. L. Sykes, Susannah Thackray-Nocera & C. Wright

East Kent Hospitals University NHS Foundation Trust, Canterbury, UK

Liam Austin, Eva Beranova, Tracey Cosier, Joanne Deery, Tracy Hazelton, Carly Price, Hazel Ramos, Reanne Solly, Sharon Turney & Heather Weston

Baillie Gifford Pandemic Science Hub, Centre for Inflammation Research, The Queen’s Medical Research Institute, University of Edinburgh, Edinburgh, UK

Nikos Avramidis, J. Kenneth Baillie, Erola Pairo-Castineira & Konrad Rawlik

Roslin Institute, University of Edinburgh, Edinburgh, UK

Nikos Avramidis, J. Kenneth Baillie & Erola Pairo-Castineira

Newcastle upon Tyne Hospitals NHS Foundation Trust and University of Newcastle, Newcastle upon Tyne, UK

A. Ayoub, J. Brown, G. Burns, Gareth Davies, Anthony De Soyza, Carlos Echevarria, Helen Fisher, C. Francis, Alan Greenhalgh, Philip Hogarth, Joan Hughes, Kasim Jiwa, G. Jones, G. MacGowan, D. Price, Avan Sayer, John Simpson, H. Tedd, S. Thomas, Sophie West, M. Witham, S. Wright & A. Young

East Cheshire NHS Trust, Macclesfield, UK

Marta Babores, Maureen Holland, Natalie Keenan, Sharlene Shashaa & Helen Wassall

Sheffield Teaching NHS Foundation Trust and University of Sheffield, Sheffield, UK

J. Bagshaw, M. Begum, K. Birchall, Robyn Butcher, H. Carborn, Flora Chan, Kerry Chapman, Yutung Cheng, Luke Chetham, Cameron Clark, Zach Coburn, Joby Cole, Myles Dixon, Alexandra Fairman, J. Finnigan, H. Foot, David Foote, Amber Ford, Rebecca Gregory, Kate Harrington, L. Haslam, L. Hesselden, J. Hockridge, Ailsa Holbourn, B. Holroyd-Hind, L. Holt, Alice Howell, E. Hurditch, F. Ilyas, Claire Jarman, Allan Lawrie, Ju Hee Lee, Elvina Lee, Rebecca Lenagh, Alison Lye, Irene Macharia, M. Marshall, Angeline Mbuyisa, J. McNeill, Sharon Megson, J. Meiring, L. Milner, S. Misra, Helen Newell, Tom Newman, C. Norman, Lorenza Nwafor, Dibya Pattenadk, Megan Plowright, Julie Porter, Phillip Ravencroft, C. Roddis, J. Rodger, Peter Saunders, J. Sidebottom, Jacqui Smith, Laurie Smith, N. Steele, G. Stephens, R. Stimpson, B. Thamu, N. Tinker, Kim Turner, Helena Turton, Phillip Wade, S. Walker, James Watson, Imogen Wilson & Amira Zawia

University of Nottingham, Nottingham, UK

David Baguley, Chris Coleman, E. Cox, Laura Fabbri, Susan Francis, Ian Hall, E. Hufton, Simon Johnson, Fasih Khan, Paaig Kitterick, Richard Morriss, Nick Selby, Iain Stewart & Louise Wright

Wirral University Teaching Hospital, Wirral, UK

Elisabeth Bailey, Anne Reddington & Andrew Wight

MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, UK

University of Swansea, Swansea, UK

University of Southampton, London, UK

David Baldwin, P. C. Calder, Nathan Huneke & Gemma Simons

Royal Brompton and Harefield Clinical Group, Guy’s and St Thomas’ NHS Foundation Trust, London, UK

R. E. Barker, Daniele Cristiano, N. Dormand, P. George, Mahitha Gummadi, S. Kon, Kamal Liyanage, C. M. Nolan, B. Patel, Suhani Patel, Oliver Polgar, L. Price, P. Shah, Suver Singh & J. A. Walsh

York and Scarborough NHS Foundation Trust, York, UK

Laura Barman, Claire Brookes, K. Elliott, L. Griffiths, Zoe Guy, Kate Howard, Diana Ionita, Heidi Redfearn, Carol Sarginson & Alison Turnbull

NHS Highland, Inverness, UK

Fiona Barrett, A. Donaldson & Beth Sage

Royal Papworth Hospital NHS Foundation Trust, Cambridge, UK

Helen Baxendale, Lucie Garner, C. Johnson, J. Mackie, Alice Michael, J. Newman, Jamie Pack, K. Paques, H. Parfrey, J. Parmar & A. Reddy

University Hospitals of Derby and Burton, Derby, UK

Paul Beckett, Caroline Dickens & Uttam Nanda

NHS Lanarkshire, Hamilton, UK

Murdina Bell, Angela Brown, M. Brown, R. Hamil, Karen Leitch, L. Macliver, Manish Patel, Jackie Quigley, Andrew Smith & B. Welsh

Cambridge University Hospitals NHS Foundation Trust, NIHR Cambridge Clinical Research Facility and University of Cambridge, Cambridge, UK

Areti Bermperi, Isabel Cruz, K. Dempsey, Anne Elmer, Jonathon Fuld, H. Jones, Sherly Jose, Stefan Marciniak, M. Parkes, Carla Ribeiro, Jessica Taylor, Mark Toshner, L. Watson & J. Worsley

Loughborough University, Loughborough, UK

Lettie Bishop & David Stensel

Betsi Cadwallader University Health Board, Bangor, UK

Annette Bolger, Ffyon Davies, Ahmed Haggar, Joanne Lewis, Arwel Lloyd, R. Manley, Emma McIvor, Daniel Menzies, K. Roberts, W. Saxon, David Southern, Christian Subbe & Victoria Whitehead

Nottingham University Hospitals NHS Trust and University of Nottingham, Nottingham, UK

Charlotte Bolton, J. Bonnington, Melanie Chrystal, Catherine Dupont, Paul Greenhaff, Ayushman Gupta, W. Jang, S. Linford, Laura Matthews, Athanasios Nikolaidis, Sabrina Prosper & Andrew Thomas

King’s College London, London, UK

Kate Bramham, M. Brown, Khalida Ismail, Tim Nicholson, Carmen Pariante, Claire Sharpe, Simon Wessely & J. Whitney

Bradford Teaching Hospitals NHS Foundation Trust, Bradford, UK

Lucy Brear, Karen Regan, Dinesh Saralaya & Kim Storton

South London and Maudsley NHS Foundation Trust and King’s College London, London, UK

G. Breen & M. Hotopf

London School of Hygiene and Tropical Medicine, London, UK

Andrew Briggs

Whittington Health NHS Trust, London, UK

E. Bright, P. Crisp, Ruvini Dharmagunawardena & M. Stern

Cardiff and Vale University Health Board, Cardiff, UK

Lauren Broad, Teriann Evans, Matthew Haynes, L. Jones, Lucy Knibbs, Alison McQueen, Catherine Oliver, Kerry Paradowski, Ramsey Sabit & Jenny Williams

Yeovil District Hospital NHS Foundation Trust, Yeovil, UK

Andrew Broadley

University of Birmingham, Birmingham, UK

Mattew Broome, Paul McArdle, Paul Moss, David Thickett, Rachel Upthegrove, Dan Wilkinson, David Wraith & Erin L. Aldera

BHF Centre for Cardiovascular Science, University of Edinburgh, Edinburgh, UK

Anda Bularga

University of Cambridge, Cambridge, UK

Ed Bullmore, Jonathon Heeney, Claudia Langenberg, William Schwaeble, Charlotte Summers & J. Weir McCall

NIHR Leicester Biomedical Research Centre–Respiratory Patient and Public Involvement Group, Leicester, UK

Jenny Bunker, Rhyan Gill & Rashmita Nathu

Imperial College Healthcare NHS Trust and Imperial College London, London, UK

L. Burden, Ellen Calvelo, Bethany Card, Caitlin Carr, Edwin Chilvers, Donna Copeland, P. Cullinan, Patrick Daly, Lynsey Evison, Tamanah Fayzan, Hussain Gordon, Sulaimaan Haq, Gisli Jenkins, Clara King, Onn Min Kon, Katherine March, Myril Mariveles, Laura McLeavey, Silvia Moriera, Unber Munawar, Uchechi Nwanguma, Lorna Orriss-Dib, Alexandra Ross, Maura Roy, Emily Russell, Katherine Samuel, J. Schronce, Neil Simpson, Lawrence Tarusan, David Thomas, Chloe Wood & Najira Yasmin

Harrogate and District NHD Foundation Trust, Harrogate, UK

Tracy Burdett, James Featherstone, Cathy Lawson, Alison Layton, Clare Mills & Lorraine Stephenson

Newcastle University/Chair of NIHR Dementia TRC, Newcastle, UK

Oxford University Hospitals NHS Foundation Trust, Oxford, UK

A. Burns & N. Kanellakis

Tameside and Glossop Integrated Care NHS Foundation Trust, Ashton-under-Lyne, UK

Al-Tahoor Butt, Martina Coulding, Heather Jones, Susan Kilroy, Jacqueline McCormick, Jerome McIntosh, Heather Savill, Victoria Turner & Joanne Vere

University of Oxford, Nuffield Department of Medicine, Oxford, UK

University of Glasgow, Glasgow, UK

Jonathon Cavanagh, S. MacDonald, Kate O’Donnell, John Petrie, Naveed Sattar & Mark Spears

United Lincolnshire Hospitals NHS Trust, Grantham, UK

Manish Chablani & Lynn Osborne

Department of Psychological Medicine, Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK

Trudie Chalder

University Hospital of South Manchester NHS Foundation Trust, Manchester, UK

N. Chaudhuri

University Hospital Southampton NHS Foundation Trust and University of Southampton, Southampton, UK

Caroline Childs, R. Djukanovic, S. Fletcher, Matt Harvey, Mark Jones, Elizabeth Marouzet, B. Marshall, Reena Samuel, T. Sass, Tim Wallis & Helen Wheeler

King’s College Hospital/Guy’s and St Thomas’ NHS FT, London, UK

A. Chiribiri & C. O’Brien

Barts Health NHS Trust, London, UK

K. Chong-James, C. David, W. Y. James, Paul Pfeffer & O. Zongo

NHS Lothian and University of Edinburgh, Edinburgh, UK

Gaunab Choudhury, S. Clohisey, Andrew Deans, J. Furniss, Ewen Harrison, S. Kelly & Aziz Sheikh

School of Cardiovascular Medicine and Sciences. King’s College London, London, UK

Phillip Chowienczyk

Lewisham and Greenwich NHS Trust, London, UK

Hywel Dda University Health Board, Haverfordwest, UK

S. Coetzee, Kim Davies, Rachel Ann Hughes, Ronda Loosley, Heather McGuinness, Abdelrahman Mohamed, Linda O’Brien, Zohra Omar, Emma Perkins, Janet Phipps, Gavin Ross, Abigail Taylor, Helen Tench & Rebecca Wolf-Roberts

NHS Tayside and University of Dundee, Dundee, UK

David Connell, C. Deas, Anne Elliott, J. George, S. Mohammed, J. Rowland, A. R. Solstice, Debbie Sutherland & Caroline Tee

Swansea Bay University Health Board, Port Talbot, UK

Lynda Connor, Amanda Cook, Gwyneth Davies, Tabitha Rees, Favas Thaivalappil & Caradog Thomas

Faculty of Medicine, Nursing and Health Sciences, School of Biomedical Sciences, Monash University, Melbourne, Victoria, Australia

Eamon Coughlan

Rotherham NHS Foundation Trust, Rotherham, UK

Alison Daniels, Anil Hormis, Julie Ingham & Lisa Zeidan

Salford Royal NHS Foundation Trust, Salford, UK

P. Dark, Nawar Diar-Bakerly, D. Evans, E. Hardy, Alice Harvey, D. Holgate, Sean Knight, N. Mairs, N. Majeed, L. McMorrow, J. Oxton, Jessica Pendlebury, C. Summersgill, R. Ugwuoke & S. Whittaker

Cwm Taf Morgannwg University Health Board, Mountain Ash, UK

Ellie Davies, Cerys Evenden, Alyson Hancock, Kia Hancock, Ceri Lynch, Meryl Rees, Lisa Roche, Natalie Stroud & T. Thomas-Woods

Borders General Hospital, NHS Borders, Melrose, UK

Joy Dawson, Hosni El-Taweel & Leanne Robinson

Aneurin Bevan University Health Board, Caerleon, UK

Amanda Dell, Sara Fairbairn, Nancy Hawkings, Jill Haworth, Michaela Hoare, Victoria Lewis, Alice Lucey, Georgia Mallison, Heeah Nassa, Chris Pennington, Andrea Price, Claire Price, Andrew Storrie, Gemma Willis & Susan Young

University of Exeter Medical School, Exeter, UK

London North West University Healthcare NHS Trust, London, UK

Shalin Diwanji, Sambasivarao Gurram, Padmasayee Papineni, Sheena Quaid, Gerlynn Tiongson & Ekaterina Watson

Alzheimer’s Research UK, Cambridge, UK

Hannah Dobson

Health and Care Research Wales, Cardiff, UK

Yvette Ellis

University of Bristol, Bristol, UK

Jonathon Evans

University of Sheffield, Sheffield, UK

L. Finnigan, Laura Saunders & James Wild

Great Western Hospital Foundation Trust, Swindon, UK

Eva Fraile & Jacinta Ugoji

Royal Devon and Exeter NHS Trust, Barnstaple, UK

Michael Gibbons

Kettering General Hospital NHS Trust, Kettering, UK

Anne-Marie Guerdette, Melanie Hewitt, R. Reddy, Katie Warwick & Sonia White

NIHR Leicester Biomedical Research Centre, Leicester, UK

Beatriz Guillen-Guio

University of Leeds, Leeds, UK

Elspeth Guthrie & Max Henderson

Royal Surrey NHS Foundation Trust, Cranleigh, UK

Mark Halling-Brown & Katherine McCullough

Chesterfield Royal Hospital NHS Trust, Calow, UK

Edward Harris & Claire Sampson

Long Covid Support, London, UK

Claire Hastie, Natalie Rogers & Nikki Smith

King’s College Hospital, NHS Foundation Trust and King’s College London, London, UK

Department of Oncology and Metabolism, University of Sheffield, Sheffield, UK

Simon Heller

NIHR Office for Clinical Research Infrastructure, London, UK

Katie Holmes

Asthma UK and British Lung Foundation Partnership, London, UK

Ian Jarrold & Samantha Walker

North Middlesex University Hospital NHS Trust, London, UK

Bhagy Jayaraman & Tessa Light

Action for Pulmonary Fibrosis, Peterborough, UK

Cardiff University, National Centre for Mental Health, Cardiff, UK

McPin Foundation, London, UK

Thomas Kabir

Roslin Institute, The University of Edinburgh, Edinburgh, UK

Steven Kerr

The Hillingdon Hospitals NHS Foundation Trust, London, UK

Samantha Kon, G. Landers, Harpreet Lota, Mariam Nasseri & Sofiya Portukhay

Queen Mary University of London, London, UK

Ania Korszun

Swansea University, Swansea Welsh Network, Hywel Dda University Health Board, Swansea, UK

Royal Infirmary of Edinburgh, NHS Lothian, Edinburgh, UK

Nazir I. Lone

Barts Heart Centre, London, UK

Barts Health NHS Trust and Queen Mary University of London, London, UK

Adrian Martineau

Salisbury NHS Foundation Trust, Salisbury, UK

Wadzanai Matimba-Mupaya & Sophia Strong-Sheldrake

University of Newcastle, Newcastle, UK

Hamish McAllister-Williams, Stella-Maria Paddick, Anthony Rostron & John Paul Taylor

Gateshead NHS Trust, Gateshead, UK

W. McCormick, Lorraine Pearce, S. Pugmire, Wendy Stoker & Ann Wilson

Manchester Centre for Clinical Neurosciences, Salford Royal NHS Foundation Trust, Manchester, UK

Katherine McIvor

Kidney Research UK, Peterborough, UK

Aisling McMahon

NHS Dumfries and Galloway, Dumfries, UK

Michael McMahon & Paula Neill

Swansea University, Swansea, UK

MQ Mental Health Research, London, UK

Lea Milligan

BHF Centre for Cardiovascular Science, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, UK

Nicholas Mills

Shropshire Community Health NHS Trust, Shropshire, UK

Sharon Painter, Johanne Tomlinson & Louise Warburton

Somerset NHS Foundation Trust, Taunton, UK

Sue Palmer, Dawn Redwood, Jo Tilley, Carinna Vickers & Tania Wainwright

Francis Crick Institute, London, UK

Markus Ralser

Manchester University NHD Foundation Trust, Manchester, UK

Pilar Rivera-Ortega

Diabetes UK, University of Glasgow, Glasgow, UK

Elizabeth Robertson

Barnsley Hospital NHS Foundation Trust, Barnsley, UK

Amy Sanderson

MRC–University of Glasgow Centre for Virus Research, Glasgow, UK

Janet Scott

Diabetes UK, London, UK

Kamini Shah

British Heart Foundation Centre, King’s College London, London, UK

King’s College Hospital NHS Foundation Trust, London, UK

University Hospitals Birmingham NHS Foundation Trust and University of Birmingham, Birmingham, UK

Institute of Cardiovascular and Medical Sciences, BHF Glasgow Cardiovascular Research Centre, University of Glasgow, Glasgow, UK

University College London NHS Foundation Trust, London and Barts Health NHS Trust, London, UK

Northumbria University, Newcastle upon Tyne, UK

Ioannis Vogiatzis

Swansea University and Swansea Welsh Network, Swansea, UK

N. Williams

DUK | NHS Digital, Salford Royal Foundation Trust, Salford, UK

Queen Alexandra Hospital, Portsmouth, UK

• Kayode Adeniji

Princess Royal Hospital, Haywards Heath, UK

Daniel Agranoff & Chi Eziefula

Bassetlaw Hospital, Bassetlaw, UK

Darent Valley Hospital, Dartford, UK

Queen Elizabeth the Queen Mother Hospital, Margate, UK

Ana Alegria

School of Informatics, University of Edinburgh, Edinburgh, UK

Beatrice Alex, Benjamin Bach & James Scott-Brown

North East and North Cumbria Ingerated, Newcastle upon Tyne, UK

Section of Biomolecular Medicine, Division of Systems Medicine, Department of Metabolism, Digestion and Reproduction, Imperial College London, London, UK

Petros Andrikopoulos, Kanta Chechi, Marc-Emmanuel Dumas, Julian Griffin, Sonia Liggi & Zoltan Takats

Section of Genomic and Environmental Medicine, Respiratory Division, National Heart and Lung Institute, Imperial College London, London, UK

Petros Andrikopoulos, Marc-Emmanuel Dumas, Michael Olanipekun & Anthonia Osagie

John Radcliffe Hospital, Oxford, UK

Brian Angus

Royal Albert Edward Infirmary, Wigan, UK

Abdul Ashish

Manchester Royal Infirmary, Manchester, UK

Dougal Atkinson

MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Crewe Road, Edinburgh, UK

Section of Molecular Virology, Imperial College London, London, UK

Wendy S. Barclay

Furness General Hospital, Barrow-in-Furness, UK

Shahedal Bari

Hull University Teaching Hospital Trust, Kingston upon Hull, UK

Gavin Barlow

Hillingdon Hospital, Hillingdon, UK

Stella Barnass

St Thomas’ Hospital, London, UK

Nicholas Barrett

Coventry and Warwickshire, Coventry, UK

Christopher Bassford

St Michael’s Hospital, Bristol, UK

Sneha Basude

Stepping Hill Hospital, Stockport, UK

David Baxter

Royal Liverpool University Hospital, Liverpool, UK

Michael Beadsworth

Bristol Royal Hospital Children’s, Bristol, UK

Jolanta Bernatoniene

Scarborough Hospital, Scarborough, UK

John Berridge

Golden Jubilee National Hospital, Clydebank, UK

Colin Berry

Liverpool Heart and Chest Hospital, Liverpool, UK

Nicola Best

Centre for Inflammation Research, The Queen’s Medical Research Institute, University of Edinburgh, Edinburgh, UK

Debby Bogaert & Clark D. Russell

James Paget University Hospital, Great Yarmouth, UK

Pieter Bothma & Darell Tupper-Carey

Aberdeen Royal Infirmary, Aberdeen, UK

Robin Brittain-Long

Adamson Hospital, Cupar, UK

Naomi Bulteel

Royal Devon and Exeter Hospital, Exeter, UK

Worcestershire Royal Hospital, Worcester, UK

Andrew Burtenshaw

ISARIC Global Support Centre, Centre for Tropical Medicine and Global Health, Nuffield Department of Medicine, University of Oxford, Oxford, UK

Gail Carson, Laura Merson & Louise Sigfrid

Conquest Hospital, Hastings, UK

Vikki Caruth

The James Cook University Hospital, Middlesbrough, UK

David Chadwick

Dorset County Hospital, Dorchester, UK

Duncan Chambler

Antimicrobial Resistance and Hospital Acquired Infection Department, Public Health England, London, UK

Meera Chand

Department of Epidemiology and Biostatistics, School of Public Health, Faculty of Medicine, Imperial College London, London, UK

Kanta Chechi

Royal Bournemouth General Hospital, Bournemouth, UK

Harrogate Hospital, Harrogate, UK

Jenny Child

Royal Blackburn Teaching Hospital, Blackburn, UK

Srikanth Chukkambotla

Edinburgh Clinical Research Facility, University of Edinburgh, Edinburgh, UK

Richard Clark, Audrey Coutts, Lorna Donelly, Angie Fawkes, Tammy Gilchrist, Katarzyna Hafezi, Louise MacGillivray, Alan Maclean, Sarah McCafferty, Kirstie Morrice, Lee Murphy & Nicola Wrobel

Torbay Hospital, Torquay, UK

Northern General Hospital, Sheffield, UK

Paul Collini, Cariad Evans & Gary Mills

Liverpool Clinical Trials Centre, University of Liverpool, Liverpool, UK

Marie Connor, Jo Dalton, Chloe Donohue, Carrol Gamble, Michelle Girvan, Sophie Halpin, Janet Harrison, Clare Jackson, Laura Marsh, Stephanie Roberts & Egle Saviciute

Department of Infectious Disease, Imperial College London, London, UK

Graham S. Cooke & Shiranee Sriskandan

St Georges Hospital (Tooting), London, UK

Catherine Cosgrove

Blackpool Victoria Hospital, Blackpool, UK

Jason Cupitt & Joanne Howard

The Royal London Hospital, London, UK

Maria-Teresa Cutino-Moguel

MRC-University of Glasgow Centre for Virus Research, Glasgow, UK

Ana da Silva Filipe, Antonia Y. W. Ho, Sarah E. McDonald, Massimo Palmarini, David L. Robertson, Janet T. Scott & Emma C. Thomson

Salford Royal Hospital, Salford, UK

University Hospital of North Durham, Durham, UK

Chris Dawson

Norfolk and Norwich University Hospital, Norwich, UK

Samir Dervisevic

Intensive Care Unit, Royal Infirmary Edinburgh, Edinburgh, UK

Annemarie B. Docherty & Seán Keating

Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool, UK

Cara Donegan & Rebecca G. Spencer

Salisbury District Hospital, Salisbury, UK

Phil Donnison

National Phenome Centre, Department of Metabolism, Digestion and Reproduction, Imperial College London, London, UK

Gonçalo dos Santos Correia, Matthew Lewis, Lynn Maslen, Caroline Sands, Zoltan Takats & Panteleimon Takis

Section of Bioanalytical Chemistry, Department of Metabolism, Digestion and Reproduction, Imperial College London, London, UK

Gonçalo dos Santos Correia, Matthew Lewis, Lynn Maslen, Caroline Sands & Panteleimon Takis

Guy’s and St Thomas’, NHS Foundation Trust, London, UK

Sam Douthwaite, Michael MacMahon, Marlies Ostermann & Manu Shankar-Hari

The Royal Oldham Hospital, Oldham, UK

Andrew Drummond

European Genomic Institute for Diabetes, Institut Pasteur de Lille, Lille University Hospital, University of Lille, Lille, France

Marc-Emmanuel Dumas

McGill University and Genome Quebec Innovation Centre, Montreal, Qeubec, Canada

National Infection Service, Public Health England, London, UK

Jake Dunning & Maria Zambon

Hereford Count Hospital, Hereford, UK

Ingrid DuRand

Southampton General Hospital, Southampton, UK

Ahilanadan Dushianthan

Northampton General Hospital, Northampton, UK

Tristan Dyer

University Hospital of Wales, Cardiff, UK

Chrisopher Fegan

University Hospitals Bristol NHS Foundation Trust, Bristol, UK

Liverpool School of Tropical Medicine, Liverpool, UK

Tom Fletcher

Leighton Hospital, Crewe, UK

Duncan Fullerton & Elijah Matovu

Manor Hospital, Walsall, UK

Scunthorpe Hospital, Scunthorpe, UK

Sanjeev Garg

Cambridge University Hospital, Cambridge, UK

Effrossyni Gkrania-Klotsas

West Suffolk NHS Foundation Trust, Bury St Edmunds, UK

Basingstoke and North Hampshire Hospital, Basingstoke, UK

Arthur Goldsmith

North Cumberland Infirmary, Carlisle, UK

Clive Graham

Paediatric Liver, GI and Nutrition Centre and MowatLabs, King’s College Hospital, London, UK

Tassos Grammatikopoulos

Institute of Liver Studies, King’s College London, London, UK

Institute of Microbiology and Infection, University of Birmingham, Birmingham, UK

Christopher A. Green

Department of Molecular and Clinical Cancer Medicine, University of Liverpool, Liverpool, UK

William Greenhalf

Institute for Global Health, University College London, London, UK

Rishi K. Gupta

NIHR Health Protection Research Unit, Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool, UK

Hayley Hardwick, Malcolm G. Semple, Tom Solomon & Lance C. W. Turtle

Warwick Hospital, Warwick, UK

Elaine Hardy

Birmingham Children’s Hospital, Birmingham, UK

Stuart Hartshorn

Nottingham City Hospital, Nottingham, UK

Daniel Harvey

Glangwili Hospital Child Health Section, Carmarthen, UK

Peter Havalda

Alder Hey Children’s Hospital, Liverpool, UK

Daniel B. Hawcutt

Department of Infectious Diseases, Queen Elizabeth University Hospital, Glasgow, UK

Antonia Y. W. Ho

Bronglais General Hospital, Aberystwyth, UK

Maria Hobrok

Worthing Hospital, Worthing, UK

Luke Hodgson

Centre for Tropical Medicine and Global Health, Nuffield Department of Medicine, University of Oxford, Oxford, UK

Peter W. Horby

Rotheram District General Hospital, Rotheram, UK

Anil Hormis

Virology Reference Department, National Infection Service, Public Health England, Colindale Avenue, London, UK

Samreen Ijaz

Royal Free Hospital, London, UK

Michael Jacobs & Padmasayee Papineni

Homerton Hospital, London, UK

Airedale Hospital, Airedale, UK

Paul Jennings

Basildon Hospital, Basildon, UK

Agilan Kaliappan

The Christie NHS Foundation Trust, Manchester, UK

Vidya Kasipandian

University Hospital Lewisham, London, UK

Stephen Kegg

The Whittington Hospital, London, UK

Michael Kelsey

Southmead Hospital, Bristol, UK

Jason Kendall

Sheffield Childrens Hospital, Sheffield, UK

Caroline Kerrison

Royal United Hospital, Bath, UK

Ian Kerslake

Department of Pharmacology, University of Liverpool, Liverpool, UK

Nuffield Department of Medicine, Peter Medawar Building for Pathogen Research, University of Oxford, Oxford, UK

Paul Klenerman

Translational Gastroenterology Unit, Nuffield Department of Medicine, University of Oxford, Oxford, UK

Public Health Scotland, Edinburgh, UK

Susan Knight, Eva Lahnsteiner & Sarah Tait

Western General Hospital, Edinburgh, UK

Oliver Koch

Southend University Hospital NHS Foundation Trust, Southend-on-Sea, UK

Gouri Koduri

Hinchingbrooke Hospital, Huntingdon, UK

George Koshy & Tamas Leiner

Royal Preston Hospital, Fulwood, UK

Shondipon Laha

University Hospital (Coventry), Coventry, UK

Steven Laird

The Walton Centre, Liverpool, UK

Susan Larkin

ISARIC, Global Support Centre, COVID-19 Clinical Research Resources, Epidemic diseases Research Group, Oxford (ERGO), University of Oxford, Oxford, UK

James Lee & Daniel Plotkin

Centre for Health Informatics, Division of Informatics, Imaging and Data Science, School of Health Sciences, Faculty of Biology, Medicine and Health, University of Manchester, Manchester Academic Health Science Centre, Manchester, UK

Gary Leeming

Hull Royal Infirmary, Hull, UK

Patrick Lillie

Nottingham University Hospitals NHS Trust:, Nottingham, UK

Wei Shen Lim

Darlington Memorial Hospital, Darlington, UK

Queen Elizabeth Hospital (Gateshead), Gateshead, UK

Vanessa Linnett

Warrington Hospital, Warrington, UK

Jeff Little

Bristol Royal Hospital for Children, Bristol, UK

Mark Lyttle

St Mary’s Hospital (Isle of Wight), Isle of Wight, UK

Emily MacNaughton

The Tunbridge Wells Hospital, Royal Tunbridge Wells, UK

Ravish Mankregod

Huddersfield Royal, Huddersfield, UK

Countess of Chester Hospital, Liverpool, UK

Ruth McEwen & Lawrence Wilson

Frimley Park Hospital, Frimley, UK

Manjula Meda

Nuffield Department of Medicine, John Radcliffe Hospital, Oxford, UK

Alexander J. Mentzer

Department of Microbiology/Infectious Diseases, Oxford University Hospitals NHS Foundation Trust, John Radcliffe Hospital, Oxford, UK

MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK

Alison M. Meynert & Murray Wham

St James University Hospital, Leeds, UK

Jane Minton

Arrowe Park Hospital, Birkenhead, UK

Kavya Mohandas

Great Ormond Street Hospital, London, UK

Royal Shrewsbury Hospital, Shrewsbury, UK

Addenbrookes Hospital, Cambridge, UK

Elinoor Moore

Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool, UK

Shona C. Moore, William A. Paxton & Georgios Pollakis

East Surrey Hospital (Redhill), Redhill, UK

Patrick Morgan

Burton Hospital, Burton, UK

Craig Morris & Tim Reynolds

Peterborough City Hospital, Peterborough, UK

Katherine Mortimore

Kent and Canterbury Hospital, Canterbury, UK

Samuel Moses

Weston Area General Trust, Bristol, UK

Mbiye Mpenge

Bedfordshire Hospital, Bedfordshire, UK

Rohinton Mulla

Glasgow Royal Infirmary, Glasgow, UK

Michael Murphy

Macclesfield General Hospital, Macclesfield, UK

Thapas Nagarajan

Derbyshire Healthcare, Derbyshire, UK

Megan Nagel

Chelsea and Westminster Hospital, London, UK

Mark Nelson & Matthew K. O’Shea

Watford General Hospital, Watford, UK

Lillian Norris & Tom Stambach

EPCC, University of Edinburgh, Edinburgh, UK

Lucy Norris

Section of Biomolecular Medicine, Division of Systems Medicine, Department of Metabolism, Digestion and Reproduction, London, UK

Michael Olanipekun

Imperial College Healthcare NHS Trust: London, London, UK

Peter J. M. Openshaw

Division of Systems Medicine, Department of Metabolism, Digestion and Reproduction, Imperial College London, London, UK

Anthonia Osagie

Prince Philip Hospital, Llanelli, UK

Igor Otahal & Andrew Workman

George Eliot Hospital – Acute Services, Nuneaton, UK

Molecular and Clinical Cancer Medicine, Institute of Systems, Molecular and Integrative Biology, University of Liverpool, Liverpool, UK

Carlo Palmieri

Clatterbridge Cancer Centre NHS Foundation Trust, Liverpool, UK

Kettering General Hospital, Kettering, UK

Selva Panchatsharam

University Hospitals of North Midlands NHS Trust, North Midlands, UK

Danai Papakonstantinou

Russells Hall Hospital, Dudley, UK

Hassan Paraiso

Harefield Hospital, Harefield, UK

Lister Hospital, Lister, UK

Natalie Pattison

Musgrove Park Hospital, Taunton, UK

Justin Pepperell

Kingston Hospital, Kingston, UK

Mark Peters

Queen’s Hospital, Romford, UK

Mandeep Phull

Southport and Formby District General Hospital, Southport, UK

Stefania Pintus

St George’s University of London, London, UK

Tim Planche

King’s College Hospital (Denmark Hill), London, UK

Centre for Clinical Infection and Diagnostics Research, Department of Infectious Diseases, School of Immunology and Microbial Sciences, King’s College London, London, UK

Nicholas Price

Department of Infectious Diseases, Guy’s and St Thomas’ NHS Foundation Trust, London, UK

The Clatterbridge Cancer Centre NHS Foundation, Bebington, UK

David Price

The Great Western Hospital, Swindon, UK

Rachel Prout

Ninewells Hospital, Dundee, UK

Nikolas Rae

Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, UK

Andrew Rambaut

Poole Hospital NHS Trust, Poole, UK

Henrik Reschreiter

William Harvey Hospital, Ashford, UK

Neil Richardson

King’s Mill Hospital, Sutton-in-Ashfield, UK

Mark Roberts

Liverpool Women’s Hospital, Liverpool, UK

Devender Roberts

Pinderfields Hospital, Wakefield, UK

Alistair Rose

North Devon District Hospital, Barnstaple, UK

Guy Rousseau

Queen Elizabeth Hospital, Birmingham, UK

Tameside General Hospital, Ashton-under-Lyne, UK

Brendan Ryan

City Hospital (Birmingham), Birmingham, UK

Taranprit Saluja

Department of Pediatrics and Virology, St Mary’s Medical School Bldg, Imperial College London, London, UK

Vanessa Sancho-Shimizu

The Newcastle Upon Tyne Hospitals NHS Foundation Trust, Newcastle Upon Tyne, UK

Matthias Schmid

NHS Greater Glasgow and Clyde, Glasgow, UK

Janet T. Scott

Respiratory Medicine, Institute in The Park, University of Liverpool, Alder Hey Children’s Hospital, Liverpool, UK

Malcolm G. Semple

Broomfield Hospital, Broomfield, UK

Stoke Mandeville, UK

Prad Shanmuga

University Hospital of North Tees, Stockton-on-Tees, UK

Anil Sharma

Institute of Translational Medicine, University of, Liverpool, Merseyside, UK

Victoria E. Shaw

Royal Manchester Children’s Hospital, Manchester, UK

Anna Shawcross

New Cross Hospital, Wolverhampton, UK

Jagtur Singh Pooni

Bedford Hospital, Bedford, UK

Jeremy Sizer

Colchester General Hospital, Colchester, UK

Richard Smith

University Hospital Birmingham NHS Foundation Trust, Birmingham, UK

Catherine Snelson & Tony Whitehouse

Walton Centre NHS Foundation Trust, Liverpool, UK

Tom Solomon

Chesterfield Royal Hospital, Calow, UK

Nick Spittle

MRC Centre for Molecular Bacteriology and Infection, Imperial College London, London, UK

Shiranee Sriskandan

Princess Alexandra Hospital, Harlow, UK

Nikki Staines & Shico Visuvanathan

Milton Keynes Hospital, Eaglestone, UK

Richard Stewart

Division of Structural Biology, The Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK

David Stuart

Royal Bolton Hopital, Farnworth, UK

Pradeep Subudhi

Department of Medicine, University of Cambridge, Cambridge, UK

Charlotte Summers

Department of Child Life and Health, University of Edinburgh, Edinburgh, UK

Olivia V. Swann

Royal Gwent (Newport), Newport, UK

Tamas Szakmany

The Royal Marsden Hospital (London), London, UK

Kate Tatham

Blood Borne Virus Unit, Virus Reference Department, National Infection Service, Public Health England, London, UK

Richard S. Tedder

Transfusion Microbiology, National Health Service Blood and Transplant, London, UK

Department of Medicine, Imperial College London, London, UK

Queen Victoria Hospital (East Grinstead), East Grinstead, UK

Leeds Teaching Hospitals NHS Trust, Leeds, UK

Robert Thompson

Royal Stoke University Hospital, Stoke-on-Trent, UK

Chris Thompson

Whiston Hospital, Rainhill, UK

Ascanio Tridente

Tropical and Infectious Disease Unit, Royal Liverpool University Hospital, Liverpool, UK

Lance C. W. Turtle

Croydon University Hospital, Thornton Heath, UK

Mary Twagira

Gloucester Royal, Gloucester, UK

Nick Vallotton

West Hertfordshire Teaching Hospitals NHS Trust, Hertfordshire, UK

Rama Vancheeswaran

North Middlesex Hospital, London, UK

Rachel Vincent

Medway Maritime Hospital, Gillingham, UK

Lisa Vincent-Smith

Royal Papworth Hospital Everard, Cambridge, UK

Alan Vuylsteke

Derriford (Plymouth), Plymouth, UK

St Helier Hospital, Sutton, UK

Rachel Wake

Royal Berkshire Hospital, Reading, UK

Andrew Walden

Royal Liverpool Hospital, Liverpool, UK

Ingeborg Welters

Bradford Royal infirmary, Bradford, UK

Paul Whittaker

Central Middlesex, London, UK

Ashley Whittington

Royal Cornwall Hospital (Tresliske), Truro, UK

Meme Wijesinghe

North Bristol NHS Trust, Bristol, UK

Martin Williams

St. Peter’s Hospital, Runnymede, UK

Stephen Winchester

Leicester Royal Infirmary, Leicester, UK

Martin Wiselka

Grantham and District Hospital, Grantham, UK

Adam Wolverson

Aintree University Hospital, Liverpool, UK

Daniel G. Wootton

North Tyneside General Hospital, North Shields, UK

Bryan Yates

Queen Elizabeth Hospital, King’s Lynn, UK

Peter Young

You can also search for this author in PubMed   Google Scholar

PHOSP-COVID collaborative group

• Kathryn Abel
• , H. Adamali
• , Davies Adeloye
• , Oluwaseun Adeyemi
• , Rita Adrego
• , Laura Aguilar Jimenez
• , Shanaz Ahmad
• , N. Ahmad Haider
• , Rubina Ahmed
• , Nyarko Ahwireng
• , Mark Ainsworth
• , Asma Alamoudi
• , Mariam Ali
• , M. Aljaroof
• , Louise Allan
• , Richard Allen
• , Lisa Allerton
• , Lynne Allsop
• , Ann Marie Allt
• , Paula Almeida
• , Bashar Al-Sheklly
• , Danny Altmann
• , Maria Alvarez Corral
• , Shannon Amoils
• , David Anderson
• , Charalambos Antoniades
• , Gill Arbane
• , Ava Maria Arias
• , Cherie Armour
• , Lisa Armstrong
• , Natalie Armstrong
• , David Arnold
• , H. Arnold
• , A. Ashish
• , Andrew Ashworth
• , M. Ashworth
• , Shahab Aslani
• , Hosanna Assefa-Kebede
• , Paul Atkin
• , Catherine Atkin
• , Raminder Aul
• , Hnin Aung
• , Liam Austin
• , Cristina Avram
• , Nikos Avramidis
• , Marta Babores
• , Rhiannon Baggott
• , J. Bagshaw
• , David Baguley
• , Elisabeth Bailey
• , J. Kenneth Baillie
• , Steve Bain
• , Majda Bakali
• , E. Baldry
• , Molly Baldwin
• , David Baldwin
• , Clive Ballard
• , Amita Banerjee
• , Dongchun Bang
• , R. E. Barker
• , Laura Barman
• , Perdita Barran
• , Shaney Barratt
• , Fiona Barrett
• , Donna Basire
• , Neil Basu
• , Michelle Bates
• , R. Batterham
• , Helen Baxendale
• , Gabrielle Baxter
• , Hannah Bayes
• , M. Beadsworth
• , Paul Beckett
• , Paul Beirne
• , Murdina Bell
• , Robert Bell
• , Kaytie Bennett
• , Eva Beranova
• , Areti Bermperi
• , Anthony Berridge
• , Colin Berry
• , Sarah Betts
• , Emily Bevan
• , Kamaldeep Bhui
• , Michelle Bingham
• , K. Birchall
• , Lettie Bishop
• , Karen Bisnauthsing
• , John Blaikely
• , Angela Bloss
• , Annette Bolger
• , Charlotte Bolton
• , J. Bonnington
• , A. Botkai
• , Charlotte Bourne
• , Michelle Bourne
• , Kate Bramham
• , Lucy Brear
• , Jonathon Breeze
• , Katie Breeze
• , Andrew Briggs
• , E. Bright
• , Christopher Brightling
• , Simon Brill
• , K. Brindle
• , Lauren Broad
• , Andrew Broadley
• , Claire Brookes
• , Mattew Broome
• , Vanessa Brown
• , Ammani Brown
• , Angela Brown
• , Jeremy Brown
• , Terry Brugha
• , Nigel Brunskill
• , Phil Buckley
• , Anda Bularga
• , Ed Bullmore
• , Jenny Bunker
• , L. Burden
• , Tracy Burdett
• , David Burn
• , John Busby
• , Robyn Butcher
• , Al-Tahoor Butt
• , P. Cairns
• , P. C. Calder
• , Ellen Calvelo
• , H. Carborn
• , Bethany Card
• , Caitlin Carr
• , Liesel Carr
• , G. Carson
• , Penny Carter
• , Anna Casey
• , M. Cassar
• , Jonathon Cavanagh
• , Manish Chablani
• , Trudie Chalder
• , James D. Chalmers
• , Rachel Chambers
• , Flora Chan
• , K. M. Channon
• , Kerry Chapman
• , Amanda Charalambou
• , N. Chaudhuri
• , A. Checkley
• , Yutung Cheng
• , Luke Chetham
• , Caroline Childs
• , Edwin Chilvers
• , H. Chinoy
• , A. Chiribiri
• , K. Chong-James
• , N. Choudhury
• , Gaunab Choudhury
• , Phillip Chowienczyk
• , C. Christie
• , Melanie Chrystal
• , Cameron Clark
• , David Clark
• , Jude Clarke
• , S. Clohisey
• , G. Coakley
• , Zach Coburn
• , S. Coetzee
• , Joby Cole
• , Chris Coleman
• , Florence Conneh
• , David Connell
• , Bronwen Connolly
• , Lynda Connor
• , Amanda Cook
• , Shirley Cooper
• , B. Cooper
• , Josh Cooper
• , Donna Copeland
• , Tracey Cosier
• , Eamon Coughlan
• , Martina Coulding
• , C. Coupland
• , Thelma Craig
• , Daniele Cristiano
• , Michael Crooks
• , Andy Cross
• , Isabel Cruz
• , P. Cullinan
• , D. Cuthbertson
• , Luke Daines
• , Matthhew Dalton
• , Patrick Daly
• , Alison Daniels
• , Joanne Dasgin
• , Anthony David
• , Ffyon Davies
• , Ellie Davies
• , Kim Davies
• , Gareth Davies
• , Gwyneth Davies
• , Melanie Davies
• , Joy Dawson
• , Camilla Dawson
• , Enya Daynes
• , Anthony De Soyza
• , Bill Deakin
• , Andrew Deans
• , Joanne Deery
• , Sylviane Defres
• , Amanda Dell
• , K. Dempsey
• , Emma Denneny
• , J. Dennis
• , Ruvini Dharmagunawardena
• , Nawar Diar-Bakerly
• , Caroline Dickens
• , A. Dipper
• , Sarah Diver
• , Shalin Diwanji
• , Myles Dixon
• , R. Djukanovic
• , Hannah Dobson
• , S. L. Dobson
• , Annemarie B. Docherty
• , A. Donaldson
• , N. Dormand
• , Andrew Dougherty
• , Rachael Dowling
• , Stephen Drain
• , Katharine Draxlbauer
• , Katie Drury
• , Pearl Dulawan
• , A. Dunleavy
• , Sarah Dunn
• , Catherine Dupont
• , Joanne Earley
• , Nicholas Easom
• , Carlos Echevarria
• , Sarah Edwards
• , C. Edwardson
• , Claudia Efstathiou
• , Anne Elliott
• , K. Elliott
• , Yvette Ellis
• , Anne Elmer
• , Omer Elneima
• , Hosni El-Taweel
• , Teriann Evans
• , Ranuromanana Evans
• , Rachael A. Evans
• , Jonathon Evans
• , Cerys Evenden
• , Lynsey Evison
• , Laura Fabbri
• , Sara Fairbairn
• , Alexandra Fairman
• , K. Fallon
• , David Faluyi
• , Clair Favager
• , Tamanah Fayzan
• , James Featherstone
• , T. Felton
• , V. Ferreira
• , Selina Finney
• , J. Finnigan
• , L. Finnigan
• , Helen Fisher
• , S. Fletcher
• , Rachel Flockton
• , Margaret Flynn
• , David Foote
• , Amber Ford
• , D. Forton
• , Eva Fraile
• , C. Francis
• , Richard Francis
• , Susan Francis
• , Anew Frankel
• , Emily Fraser
• , N. French
• , Jonathon Fuld
• , J. Furniss
• , Lucie Garner
• , N. Gautam
• , John Geddes
• , J. George
• , P. George
• , Michael Gibbons
• , Rhyan Gill
• , Mandy Gill
• , L. Gilmour
• , F. Gleeson
• , Jodie Glossop
• , Sarah Glover
• , Nicola Goodman
• , Camelia Goodwin
• , Bibek Gooptu
• , Hussain Gordon
• , T. Gorsuch
• , M. Greatorex
• , Paul Greenhaff
• , William Greenhalf
• , Alan Greenhalgh
• , Neil J. Greening
• , John Greenwood
• , Rebecca Gregory
• , Heidi Gregory
• , D. Grieve
• , Denise Griffin
• , L. Griffiths
• , Anne-Marie Guerdette
• , Beatriz Guillen-Guio
• , Mahitha Gummadi
• , Ayushman Gupta
• , Sambasivarao Gurram
• , Elspeth Guthrie
• , Kate Hadley
• , Ahmed Haggar
• , Kera Hainey
• , Brigid Hairsine
• , Pranab Haldar
• , Lucy Hall
• , Mark Halling-Brown
• , Alyson Hancock
• , Kia Hancock
• , Neil Hanley
• , Sulaimaan Haq
• , Hayley Hardwick
• , Tim Hardy
• , Beverley Hargadon
• , Kate Harrington
• , Edward Harris
• , Victoria C. Harris
• , Ewen Harrison
• , Paul Harrison
• , Nicholas Hart
• , Alice Harvey
• , Matt Harvey
• , M. Harvie
• , L. Haslam
• , Claire Hastie
• , May Havinden-Williams
• , Jenny Hawkes
• , Nancy Hawkings
• , Jill Haworth
• , A. Hayday
• , Matthew Haynes
• , J. Hazeldine
• , Tracy Hazelton
• , Liam Heaney
• , Cheryl Heeley
• , Jonathon Heeney
• , M. Heightman
• , Simon Heller
• , Max Henderson
• , Helen Henson
• , L. Hesselden
• , Melanie Hewitt
• , Victoria Highett
• , T. Hillman
• , Ling-Pei Ho
• , Michaela Hoare
• , Amy Hoare
• , J. Hockridge
• , Philip Hogarth
• , Ailsa Holbourn
• , Sophie Holden
• , L. Holdsworth
• , D. Holgate
• , Maureen Holland
• , Leah Holloway
• , Katie Holmes
• , Megan Holmes
• , B. Holroyd-Hind
• , Anil Hormis
• , Alexander Horsley
• , Akram Hosseini
• , M. Hotopf
• , Linzy Houchen-Wolloff
• , Luke S. Howard
• , Kate Howard
• , Alice Howell
• , E. Hufton
• , Rachel Ann Hughes
• , Joan Hughes
• , Alun Hughes
• , Amy Humphries
• , Nathan Huneke
• , E. Hurditch
• , John Hurst
• , Masud Husain
• , Tracy Hussell
• , John Hutchinson
• , W. Ibrahim
• , Julie Ingham
• , L. Ingram
• , Diana Ionita
• , Karen Isaacs
• , Khalida Ismail
• , T. Jackson
• , Joseph Jacob
• , W. Y. James
• , Claire Jarman
• , Ian Jarrold
• , Hannah Jarvis
• , Roman Jastrub
• , Bhagy Jayaraman
• , Gisli Jenkins
• , P. Jezzard
• , Kasim Jiwa
• , C. Johnson
• , Simon Johnson
• , Desmond Johnston
• , Caroline Jolley
• , Ian Jones
• , Heather Jones
• , Mark Jones
• , Don Jones
• , Sherly Jose
• , Thomas Kabir
• , G. Kaltsakas
• , Vicky Kamwa
• , N. Kanellakis
• , Sabina Kaprowska
• , Zunaira Kausar
• , Natalie Keenan
• , Steven Kerr
• , Helen Kerslake
• , Angela Key
• , Fasih Khan
• , Kamlesh Khunti
• , Susan Kilroy
• , Bernie King
• , Clara King
• , Lucy Kingham
• , Jill Kirk
• , Paaig Kitterick
• , Paul Klenerman
• , Lucy Knibbs
• , Sean Knight
• , Abigail Knighton
• , Onn Min Kon
• , Samantha Kon
• , Ania Korszun
• , Ivan Koychev
• , Claire Kurasz
• , Prathiba Kurupati
• , Hanan Lamlum
• , G. Landers
• , Claudia Langenberg
• , Lara Lavelle-Langham
• , Allan Lawrie
• , Cathy Lawson
• , Claire Lawson
• , Alison Layton
• , Olivia C. Leavy
• , Ju Hee Lee
• , Elvina Lee
• , Karen Leitch
• , Rebecca Lenagh
• , Victoria Lewis
• , Joanne Lewis
• , Keir Lewis
• , N. Lewis-Burke
• , Felicity Liew
• , Tessa Light
• , Liz Lightstone
• , W. Lilaonitkul
• , S. Linford
• , Anne Lingford-Hughes
• , M. Lipman
• , Kamal Liyanage
• , Arwel Lloyd
• , Nazir I. Lone
• , Ronda Loosley
• , Janet Lord
• , Harpreet Lota
• , Wayne Lovegrove
• , Daniel Lozano-Rojas
• , Alice Lucey
• , Gardiner Lucy
• , E. Lukaschuk
• , Alison Lye
• , Ceri Lynch
• , S. MacDonald
• , G. MacGowan
• , Irene Macharia
• , J. Mackie
• , L. Macliver
• , S. Madathil
• , Gladys Madzamba
• , Nick Magee
• , Murphy Magtoto
• , N. Majeed
• , Flora Malein
• , Georgia Mallison
• , William Man
• , S. Mandal
• , K. Mangion
• , C. Manisty
• , R. Manley
• , Katherine March
• , Stefan Marciniak
• , Philip Marino
• , Myril Mariveles
• , Michael Marks
• , Elizabeth Marouzet
• , Sophie Marsh
• , M. Marshall
• , B. Marshall
• , Jane Martin
• , Adrian Martineau
• , L. M. Martinez
• , Nick Maskell
• , Darwin Matila
• , Wadzanai Matimba-Mupaya
• , Laura Matthews
• , Angeline Mbuyisa
• , Steve McAdoo
• , Hamish McAllister-Williams
• , Paul McArdle
• , Anne McArdle
• , Danny McAulay
• , Hamish J. C. McAuley
• , Gerry McCann
• , W. McCormick
• , Jacqueline McCormick
• , P. McCourt
• , Celeste McCracken
• , Lorcan McGarvey
• , Jade McGinness
• , K. McGlynn
• , Andrew McGovern
• , Heather McGuinness
• , I. B. McInnes
• , Jerome McIntosh
• , Emma McIvor
• , Katherine McIvor
• , Laura McLeavey
• , Aisling McMahon
• , Michael McMahon
• , L. McMorrow
• , Teresa Mcnally
• , M. McNarry
• , J. McNeill
• , Alison McQueen
• , H. McShane
• , Chloe Mears
• , Clare Megson
• , Sharon Megson
• , J. Meiring
• , Lucy Melling
• , Mark Mencias
• , Daniel Menzies
• , Marta Merida Morillas
• , Alice Michael
• , Benedict Michael
• , C. A. Miller
• , Lea Milligan
• , Nicholas Mills
• , Clare Mills
• , George Mills
• , L. Milner
• , Jane Mitchell
• , Abdelrahman Mohamed
• , Noura Mohamed
• , S. Mohammed
• , Philip Molyneaux
• , Will Monteiro
• , Silvia Moriera
• , Anna Morley
• , Leigh Morrison
• , Richard Morriss
• , A. Morrow
• , Paul Moss
• , Alistair Moss
• , K. Motohashi
• , N. Msimanga
• , Elizabeta Mukaetova-Ladinska
• , Unber Munawar
• , Jennifer Murira
• , Uttam Nanda
• , Heeah Nassa
• , Mariam Nasseri
• , Rashmita Nathu
• , Aoife Neal
• , Robert Needham
• , Paula Neill
• , Stefan Neubauer
• , D. E. Newby
• , Helen Newell
• , J. Newman
• , Tom Newman
• , Alex Newton-Cox
• , T. E. Nichols
• , Tim Nicholson
• , Christos Nicolaou
• , Debby Nicoll
• , Athanasios Nikolaidis
• , C. Nikolaidou
• , C. M. Nolan
• , Matthew Noonan
• , C. Norman
• , Petr Novotny
• , Kimon Ntotsis
• , Jose Nunag
• , Lorenza Nwafor
• , Uchechi Nwanguma
• , Joseph Nyaboko
• , Linda O’Brien
• , C. O’Brien
• , Natasha Odell
• , Kate O’Donnell
• , Godwin Ogbole
• , Olaoluwa Olaosebikan
• , Catherine Oliver
• , Zohra Omar
• , Peter J. M. Openshaw
• , D. P. O’Regan
• , Lorna Orriss-Dib
• , Lynn Osborne
• , Rebecca Osbourne
• , Marlies Ostermann
• , Charlotte Overton
• , Jamie Pack
• , Edmund Pacpaco
• , Stella-Maria Paddick
• , Sharon Painter
• , Erola Pairo-Castineira
• , Ashkan Pakzad
• , Sue Palmer
• , Padmasayee Papineni
• , K. Paques
• , Kerry Paradowski
• , Manish Pareek
• , Dhruv Parekh
• , H. Parfrey
• , Carmen Pariante
• , S. Parker
• , M. Parkes
• , J. Parmar
• , Sheetal Patale
• , Manish Patel
• , Suhani Patel
• , Dibya Pattenadk
• , M. Pavlides
• , Sheila Payne
• , Lorraine Pearce
• , John Pearl
• , Dan Peckham
• , Jessica Pendlebury
• , Yanchun Peng
• , Chris Pennington
• , Ida Peralta
• , Emma Perkins
• , Z. Peterkin
• , Tunde Peto
• , Nayia Petousi
• , John Petrie
• , Paul Pfeffer
• , Janet Phipps
• , S. Piechnik
• , John Pimm
• , Karen Piper Hanley
• , Riinu Pius
• , Hannah Plant
• , Tatiana Plekhanova
• , Megan Plowright
• , Krisnah Poinasamy
• , Oliver Polgar
• , Julie Porter
• , Joanna Porter
• , Sofiya Portukhay
• , Natassia Powell
• , A. Prabhu
• , James Pratt
• , Andrea Price
• , Claire Price
• , Carly Price
• , Anne Prickett
• , I. Propescu
• , J. Propescu
• , Sabrina Prosper
• , S. Pugmire
• , Sheena Quaid
• , Jackie Quigley
• , Jennifer K. Quint
• , H. Qureshi
• , I. N. Qureshi
• , K. Radhakrishnan
• , Najib Rahman
• , Markus Ralser
• , Betty Raman
• , Hazel Ramos
• , Albert Ramos
• , Jade Rangeley
• , Bojidar Rangelov
• , Liz Ratcliffe
• , Phillip Ravencroft
• , Konrad Rawlik
• , Anne Reddington
• , Heidi Redfearn
• , Dawn Redwood
• , Annabel Reed
• , Meryl Rees
• , Tabitha Rees
• , Karen Regan
• , Will Reynolds
• , Carla Ribeiro
• , A. Richards
• , Emma Richardson
• , M. Richardson
• , Pilar Rivera-Ortega
• , K. Roberts
• , Elizabeth Robertson
• , Leanne Robinson
• , Emma Robinson
• , Lisa Roche
• , C. Roddis
• , J. Rodger
• , Natalie Rogers
• , Gavin Ross
• , Alexandra Ross
• , Jennifer Rossdale
• , Anthony Rostron
• , Anna Rowe
• , J. Rowland
• , M. J. Rowland
• , A. Rowland
• , Sarah L. Rowland-Jones
• , Maura Roy
• , Igor Rudan
• , Richard Russell
• , Emily Russell
• , Gwen Saalmink
• , Ramsey Sabit
• , Beth Sage
• , T. Samakomva
• , Nilesh Samani
• , A. A. Samat
• , Claire Sampson
• , Katherine Samuel
• , Reena Samuel
• , Z. B. Sanders
• , Amy Sanderson
• , Elizabeth Sapey
• , Dinesh Saralaya
• , Jack Sargant
• , Carol Sarginson
• , Naveed Sattar
• , Kathryn Saunders
• , Peter Saunders
• , Ruth Saunders
• , Laura Saunders
• , Heather Savill
• , Avan Sayer
• , J. Schronce
• , William Schwaeble
• , Janet Scott
• , Kathryn Scott
• , Nick Selby
• , Malcolm G. Semple
• , Marco Sereno
• , Terri Ann Sewell
• , Kamini Shah
• , Ajay Shah
• , Manu Shankar-Hari
• , M. Sharma
• , Claire Sharpe
• , Michael Sharpe
• , Sharlene Shashaa
• , Alison Shaw
• , Victoria Shaw
• , Karen Shaw
• , Aziz Sheikh
• , Sarah Shelton
• , Liz Shenton
• , K. Shevket
• , Aarti Shikotra
• , Sulman Siddique
• , Salman Siddiqui
• , J. Sidebottom
• , Louise Sigfrid
• , Gemma Simons
• , Neil Simpson
• , John Simpson
• , Ananga Singapuri
• , Suver Singh
• , Claire Singh
• , Sally Singh
• , D. Sissons
• , J. Skeemer
• , Katie Slack
• , David Smith
• , Nikki Smith
• , Andrew Smith
• , Jacqui Smith
• , Laurie Smith
• , Susan Smith
• , M. Soares
• , Teresa Solano
• , Reanne Solly
• , A. R. Solstice
• , Tracy Soulsby
• , David Southern
• , D. Sowter
• , Mark Spears
• , Lisa Spencer
• , Fabio Speranza
• , Louise Stadon
• , Stefan Stanel
• , R. Steeds
• , N. Steele
• , Mike Steiner
• , David Stensel
• , G. Stephens
• , Lorraine Stephenson
• , Iain Stewart
• , R. Stimpson
• , Sue Stockdale
• , J. Stockley
• , Wendy Stoker
• , Roisin Stone
• , Will Storrar
• , Andrew Storrie
• , Kim Storton
• , E. Stringer
• , Sophia Strong-Sheldrake
• , Natalie Stroud
• , Christian Subbe
• , Catherine Sudlow
• , Zehra Suleiman
• , Charlotte Summers
• , C. Summersgill
• , Debbie Sutherland
• , D. L. Sykes
• , Nick Talbot
• , Ai Lyn Tan
• , Lawrence Tarusan
• , Vera Tavoukjian
• , Jessica Taylor
• , Abigail Taylor
• , Chris Taylor
• , John Paul Taylor
• , Amelie Te
• , Caroline Tee
• , J. Teixeira
• , Helen Tench
• , Sarah Terry
• , Susannah Thackray-Nocera
• , Favas Thaivalappil
• , David Thickett
• , David Thomas
• , S. Thomas
• , Caradog Thomas
• , Andrew Thomas
• , T. Thomas-Woods
• , A. A. Roger Thompson
• , Tamika Thompson
• , T. Thornton
• , Matthew Thorpe
• , Ryan S. Thwaites
• , Jo Tilley
• , N. Tinker
• , Gerlynn Tiongson
• , Martin Tobin
• , Johanne Tomlinson
• , Mark Toshner
• , T. Treibel
• , K. A. Tripp
• , Drupad Trivedi
• , E. M. Tunnicliffe
• , Alison Turnbull
• , Kim Turner
• , Sarah Turner
• , Victoria Turner
• , E. Turner
• , Sharon Turney
• , Lance Turtle
• , Helena Turton
• , Jacinta Ugoji
• , R. Ugwuoke
• , Rachel Upthegrove
• , Jonathon Valabhji
• , Maximina Ventura
• , Joanne Vere
• , Carinna Vickers
• , Ben Vinson
• , Ioannis Vogiatzis
• , Elaine Wade
• , Phillip Wade
• , Louise V. Wain
• , Tania Wainwright
• , Lilian Wajero
• , Sinead Walder
• , Samantha Walker
• , S. Walker
• , Tim Wallis
• , Sarah Walmsley
• , Simon Walsh
• , J. A. Walsh
• , Louise Warburton
• , T. J. C. Ward
• , Katie Warwick
• , Helen Wassall
• , Samuel Waterson
• , L. Watson
• , Ekaterina Watson
• , James Watson
• , M. Webster
• , J. Weir McCall
• , Carly Welch
• , Simon Wessely
• , Sophie West
• , Heather Weston
• , Helen Wheeler
• , Sonia White
• , Victoria Whitehead
• , J. Whitney
• , S. Whittaker
• , Beverley Whittam
• , V. Whitworth
• , Andrew Wight
• , James Wild
• , Martin Wilkins
• , Dan Wilkinson
• , Nick Williams
• , N. Williams
• , B. Williams
• , Jenny Williams
• , S. A. Williams-Howard
• , Michelle Willicombe
• , Gemma Willis
• , James Willoughby
• , Ann Wilson
• , Imogen Wilson
• , Daisy Wilson
• , Nicola Window
• , M. Witham
• , Rebecca Wolf-Roberts
• , Chloe Wood
• , F. Woodhead
• , Janet Woods
• , Dan Wootton
• , J. Wormleighton
• , J. Worsley
• , David Wraith
• , Caroline Wrey Brown
• , C. Wright
• , S. Wright
• , Louise Wright
• , Inez Wynter
• , Moucheng Xu
• , Najira Yasmin
• , S. Yasmin
• , Tom Yates
• , Kay Por Yip
• , Susan Young
• , Bob Young
• , A. J. Yousuf
• , Amira Zawia
• , Lisa Zeidan
• , Bang Zhao
• , Bang Zheng
•  & O. Zongo
• , Daniel Agranoff
• , Ken Agwuh
• , Katie A. Ahmed
• , Dhiraj Ail
• , Erin L. Aldera
• , Ana Alegria
• , Beatrice Alex
• , Sam Allen
• , Petros Andrikopoulos
• , Brian Angus
• , Jane A. Armstrong
• , Abdul Ashish
• , Milton Ashworth
• , Innocent G. Asiimwe
• , Dougal Atkinson
• , Benjamin Bach
• , Siddharth Bakshi
• , Wendy S. Barclay
• , Shahedal Bari
• , Gavin Barlow
• , Samantha L. Barlow
• , Stella Barnass
• , Nicholas Barrett
• , Christopher Bassford
• , Sneha Basude
• , David Baxter
• , Michael Beadsworth
• , Jolanta Bernatoniene
• , John Berridge
• , Nicola Best
• , Debby Bogaert
• , Laura Booth
• , Pieter Bothma
• , Benjamin Brennan
• , Robin Brittain-Long
• , Katie Bullock
• , Naomi Bulteel
• , Tom Burden
• , Andrew Burtenshaw
• , Nicola Carlucci
• , Gail Carson
• , Vikki Caruth
• , Emily Cass
• , Benjamin W. A. Catterall
• , David Chadwick
• , Duncan Chambler
• , Meera Chand
• , Kanta Chechi
• , Nigel Chee
• , Jenny Child
• , Srikanth Chukkambotla
• , Richard Clark
• , Tom Clark
• , Jordan J. Clark
• , Emily A. Clarke
• , Sara Clohisey
• , Sarah Cole
• , Paul Collini
• , Marie Connor
• , Graham S. Cooke
• , Louise Cooper
• , Catherine Cosgrove
• , Audrey Coutts
• , Helen Cox
• , Jason Cupitt
• , Maria-Teresa Cutino-Moguel
• , Ana da Silva Filipe
• , Jo Dalton
• , Paul Dark
• , Christopher Davis
• , Chris Dawson
• , Thushan de Silva
• , Samir Dervisevic
• , Oslem Dincarslan
• , Alejandra Doce Carracedo
• , Cara Donegan
• , Lorna Donelly
• , Phil Donnison
• , Chloe Donohue
• , Gonçalo dos Santos Correia
• , Sam Douthwaite
• , Thomas M. Drake
• , Andrew Drummond
• , Marc-Emmanuel Dumas
• , Chris Dunn
• , Jake Dunning
• , Ingrid DuRand
• , Ahilanadan Dushianthan
• , Tristan Dyer
• , Philip Dyer
• , Angela Elliott
• , Cariad Evans
• , Anthony Evans
• , Chi Eziefula
• , Cameron J. Fairfield
• , Angie Fawkes
• , Chrisopher Fegan
• , Lorna Finch
• , Adam Finn
• , Lewis W. S. Fisher
• , Lisa Flaherty
• , Tom Fletcher
• , Terry Foster
• , Duncan Fullerton
• , Carrol Gamble
• , Isabel Garcia-Dorival
• , Atul Garg
• , Sanjeev Garg
• , Tammy Gilchrist
• , Michelle Girvan
• , Effrossyni Gkrania-Klotsas
• , Jo Godden
• , Arthur Goldsmith
• , Clive Graham
• , Tassos Grammatikopoulos
• , Christopher A. Green
• , Julian Griffin
• , Fiona Griffiths
• , Philip Gunning
• , Rishi K. Gupta
• , Katarzyna Hafezi
• , Sophie Halpin
• , Elaine Hardy
• , Ewen M. Harrison
• , Janet Harrison
• , Catherine Hartley
• , Stuart Hartshorn
• , Daniel Harvey
• , Peter Havalda
• , Daniel B. Hawcutt
• , Ross Hendry
• , Antonia Y. W. Ho
• , Maria Hobrok
• , Luke Hodgson
• , Karl Holden
• , Anthony Holmes
• , Peter W. Horby
• , Joanne Howard
• , Samreen Ijaz
• , Clare Jackson
• , Michael Jacobs
• , Susan Jain
• , Paul Jennings
• , Rebecca L. Jensen
• , Christopher B. Jones
• , Trevor R. Jones
• , Agilan Kaliappan
• , Vidya Kasipandian
• , Seán Keating
• , Stephen Kegg
• , Michael Kelsey
• , Jason Kendall
• , Caroline Kerrison
• , Ian Kerslake
• , Shadia Khandaker
• , Katharine King
• , Robyn T. Kiy
• , Stephen R. Knight
• , Susan Knight
• , Oliver Koch
• , Gouri Koduri
• , George Koshy
• , Chrysa Koukorava
• , Shondipon Laha
• , Eva Lahnsteiner
• , Steven Laird
• , Annette Lake
• , Suzannah Lant
• , Susan Larkin
• , Diane Latawiec
• , Andrew Law
• , James Lee
• , Gary Leeming
• , Daniella Lefteri
• , Tamas Leiner
• , Lauren Lett
• , Matthew Lewis
• , Sonia Liggi
• , Patrick Lillie
• , Wei Shen Lim
• , James Limb
• , Vanessa Linnett
• , Jeff Little
• , Lucia A. Livoti
• , Mark Lyttle
• , Louise MacGillivray
• , Alan Maclean
• , Michael MacMahon
• , Emily MacNaughton
• , Maria Mancini
• , Ravish Mankregod
• , Laura Marsh
• , Lynn Maslen
• , Hannah Massey
• , Huw Masson
• , Elijah Matovu
• , Nicole Maziere
• , Sarah McCafferty
• , Katherine McCullough
• , Sarah E. McDonald
• , Sarah McDonald
• , Laurence McEvoy
• , Ruth McEwen
• , John McLauchlan
• , Kenneth A. Mclean
• , Manjula Meda
• , Alexander J. Mentzer
• , Laura Merson
• , Soeren Metelmann
• , Alison M. Meynert
• , Nahida S. Miah
• , Joanna Middleton
• , Gary Mills
• , Jane Minton
• , Joyce Mitchell
• , Kavya Mohandas
• , James Moon
• , Elinoor Moore
• , Shona C. Moore
• , Patrick Morgan
• , Kirstie Morrice
• , Craig Morris
• , Katherine Mortimore
• , Samuel Moses
• , Mbiye Mpenge
• , Rohinton Mulla
• , Derek Murphy
• , Lee Murphy
• , Michael Murphy
• , Ellen G. Murphy
• , Thapas Nagarajan
• , Megan Nagel
• , Mark Nelson
• , Lisa Norman
• , Lillian Norris
• , Lucy Norris
• , Mahdad Noursadeghi
• , Michael Olanipekun
• , Wilna Oosthuyzen
• , Anthonia Osagie
• , Matthew K. O’Shea
• , Igor Otahal
• , Mark Pais
• , Massimo Palmarini
• , Carlo Palmieri
• , Selva Panchatsharam
• , Danai Papakonstantinou
• , Hassan Paraiso
• , Brij Patel
• , Natalie Pattison
• , William A. Paxton
• , Rebekah Penrice-Randal
• , Justin Pepperell
• , Mark Peters
• , Mandeep Phull
• , Jack Pilgrim
• , Stefania Pintus
• , Tim Planche
• , Daniel Plotkin
• , Georgios Pollakis
• , Frank Post
• , Nicholas Price
• , David Price
• , Tessa Prince
• , Rachel Prout
• , Nikolas Rae
• , Andrew Rambaut
• , Henrik Reschreiter
• , Tim Reynolds
• , Neil Richardson
• , P. Matthew Ridley
• , Mark Roberts
• , Stephanie Roberts
• , Devender Roberts
• , David L. Robertson
• , Alistair Rose
• , Guy Rousseau
• , Bobby Ruge
• , Clark D. Russell
• , Brendan Ryan
• , Debby Sales
• , Taranprit Saluja
• , Vanessa Sancho-Shimizu
• , Caroline Sands
• , Egle Saviciute
• , Matthias Schmid
• , Janet T. Scott
• , James Scott-Brown
• , Aarti Shah
• , Prad Shanmuga
• , Anil Sharma
• , Catherine A. Shaw
• , Victoria E. Shaw
• , Anna Shawcross
• , Rebecca K. Shears
• , Jagtur Singh Pooni
• , Jeremy Sizer
• , Benjamin Small
• , Richard Smith
• , Catherine Snelson
• , Tom Solomon
• , Rebecca G. Spencer
• , Nick Spittle
• , Shiranee Sriskandan
• , Nikki Staines
• , Tom Stambach
• , Richard Stewart
• , David Stuart
• , Krishanthi S. Subramaniam
• , Pradeep Subudhi
• , Olivia V. Swann
• , Tamas Szakmany
• , Agnieska Szemiel
• , Aislynn Taggart
• , Sarah Tait
• , Zoltan Takats
• , Panteleimon Takis
• , Jolanta Tanianis-Hughes
• , Kate Tatham
• , Richard S. Tedder
• , Jo Thomas
• , Jordan Thomas
• , Robert Thompson
• , Chris Thompson
• , Emma C. Thomson
• , Ascanio Tridente
• , Erwan Trochu
• , Darell Tupper-Carey
• , Lance C. W. Turtle
• , Mary Twagira
• , Nick Vallotton
• , Libby van Tonder
• , Rama Vancheeswaran
• , Rachel Vincent
• , Lisa Vincent-Smith
• , Shico Visuvanathan
• , Alan Vuylsteke
• , Sam Waddy
• , Rachel Wake
• , Andrew Walden
• , Ingeborg Welters
• , Murray Wham
• , Tony Whitehouse
• , Paul Whittaker
• , Ashley Whittington
• , Meme Wijesinghe
• , Eve Wilcock
• , Martin Williams
• , Lawrence Wilson
• , Stephen Winchester
• , Martin Wiselka
• , Adam Wolverson
• , Daniel G. Wootton
• , Andrew Workman
• , Nicola Wrobel
• , Bryan Yates
• , Peter Young
• , Maria Zambon
•  & J. Eunice Zhang

Contributions

F.L. recruited participants, acquired clinical samples, analyzed and interpreted data and cowrote the manuscript, including all drafting and revisions. C.E. analyzed and interpreted data and cowrote this manuscript, including all drafting and revisions. S.F. and M.R. supported the analysis and interpretation of data as well as drafting and revisions. D.S., J.K.S., S.C.M., S.A., N.M., J.N., C.K., O.C.L., O.E., H.J.C.M., A. Shikotra, A. Singapuri, M.S., V.C.H., M.T., N.J.G., N.I.L. and C.C. contributed to acquisition of data underlying this study. L.H.-W., A.A.R.T., S.L.R.-J., L.S.H., O.M.K., D.G.W., T.I.d.S. and A. Ho made substantial contributions to conception/design and implementation of this work and/or acquisition of clinical samples for this work. They have supported drafting and revisions of the manuscript. E.M.H., J.K.Q. and A.B.D. made substantial contributions to the study design as well as data access, linkage and analysis. They have supported drafting and revisions of this work. J.D.C., L.-P.H., A. Horsley, B.R., K.P., M.M. and W.G. made substantial contributions to the conception and design of this work and have supported drafting and revisions of this work. J.K.B. obtained funding for ISARIC4C, is ISARIC4C consortium co-lead, has made substantial contributions to conception and design of this work and has supported drafting and revisions of this work. M.G.S. obtained funding for ISARIC4C, is ISARIC4C consortium co-lead, sponsor/protocol chief investigator, has made substantial contributions to conception and design of this work and has supported drafting and revisions of this work. R.A.E. and L.V.W. are co-leads of PHOSP-COVID, made substantial contributions to conception and design of this work, the acquisition and analysis of data, and have supported drafting and revisions of this work. C.B. is the chief investigator of PHOSP-COVID and has made substantial contributions to conception and design of this work. R.S.T. and L.T. made substantial contributions to the acquisition, analysis and interpretation of the data underlying this study and have contributed to drafting and revisions of this work. P.J.M.O. obtained funding for ISARIC4C, is ISARIC4C consortium co-lead, sponsor/protocol chief investigator and has made substantial contributions to conception and design of this work. R.S.T. and P.J.M.O. have also made key contributions to interpretation of data and have co-written this manuscript. All authors have read and approve the final version to be published. All authors agree to accountability for all aspects of this work. All investigators within ISARIC4C and the PHOSP-COVID consortia have made substantial contributions to the conception or design of this study and/or acquisition of data for this study. The full list of authors within these groups is available in Supplementary Information .

Corresponding authors

Correspondence to Ryan S. Thwaites or Peter J. M. Openshaw .

Ethics declarations

Competing interests.

F.L., C.E., D.S., J.K.S., S.C.M., C.D., C.K., N.M., L.N., E.M.H., A.B.D., J.K.Q., L.-P.H., K.P., L.S.H., O.M.K., S.F., T.I.d.S., D.G.W., R.S.T. and J.K.B. have no conflicts of interest. A.A.R.T. receives speaker fees and support to attend meetings from Janssen Pharmaceuticals. S.L.R.-J. is on the data safety monitoring board for Bexero trial in HIV+ adults in Kenya. J.D.C. is the deputy chief editor of the European Respiratory Journal and receives consulting fees from AstraZeneca, Boehringer Ingelheim, Chiesi, GSK, Insmed, Janssen, Novartis, Pfizer and Zambon. A. Horsley is deputy chair of NIHR Translational Research Collaboration (unpaid role). B.R. receives honoraria from Axcella therapeutics. R.A.E. is co-lead of PHOSP-COVID and receives fees from AstraZenaca/Evidera for consultancy on LC and from AstraZenaca for consultancy on digital health. R.A.E. has received speaker fees from Boehringer in June 2021 and has held a role as European Respiratory Society Assembly 01.02 Pulmonary Rehabilitation secretary. R.A.E. is on the American Thoracic Society Pulmonary Rehabilitation Assembly program committee. L.V.W. also receives funding from Orion pharma and GSK and holds contracts with Genentech and AstraZenaca. L.V.W. has received consulting fees from Galapagos and Boehringer, is on the data advisory board for Galapagos and is Associate Editor for the European Respiratory Journal . A. Ho is a member of NIHR Urgent Public Health Group (June 2020–March 2021). M.M. is an applicant on the PHOSP study funded by NIHR/DHSC. M.G.S. acts as an independent external and nonremunerated member of Pfizer’s External Data Monitoring Committee for their mRNA vaccine program(s), is Chair of Infectious Disease Scientific Advisory Board of Integrum Scientific LLC, and is director of MedEx Solutions Ltd. and majority owner of MedEx Solutions Ltd. and minority owner of Integrum Scientific LLC. M.G.S.’s institution has been in receipt of gifts from Chiesi Farmaceutici S.p.A. of Clinical Trial Investigational Medicinal Product without encumbrance and distribution of same to trial sites. M.G.S. is a nonrenumerated member of HMG UK New Emerging Respiratory Virus Threats Advisory Group and has previously been a nonrenumerated member of the Scientific Advisory Group for Emergencies (SAGE). C.B. has received consulting fees and/or grants from GSK, AstraZeneca, Genentech, Roche, Novartis, Sanofi, Regeneron, Chiesi, Mologic and 4DPharma. L.T. has received consulting fees from MHRA, AstraZeneca and Synairgen and speakers’ fees from Eisai Ltd., and support for conference attendance from AstraZeneca. L.T. has a patent pending with ZikaVac. P.J.M.O. reports grants from the EU Innovative Medicines Initiative 2 Joint Undertaking during the submitted work; grants from UK Medical Research Council, GSK, Wellcome Trust, EU Innovative Medicines Initiative, UK National Institute for Health Research and UK Research and Innovation–Department for Business, Energy and Industrial Strategy; and personal fees from Pfizer, Janssen and Seqirus, outside the submitted work.

Peer review

Peer review information.

Nature Immunology thanks Ziyad Al-Aly and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Ioana Staicu was the primary editor on this article and managed its editorial process and peer review in collaboration with the rest of the editorial team.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data

Extended data fig. 1 penalized logistic regression performance..

Graphs show classification error and Area under curve (AUC) from the 50 repeats tenfold nested cross-validation used to optimise and assess the performance of PLR testing associations with each LC outcome relative to Recovered (n = 233): Cardio_Resp (n = 398), Fatigue (n = 384), Anxiety/Depression (n = 202), GI (n = 132), ( e ) Cognitive (n = 6). The distributions of classification error and area under curve (AUC) from the nested cross-validation are shown. Box plot centre line represents the Median and boundaries of the box represent interquartile range (IQR), the whisker length represent 1.5xIQR.

Extended Data Fig. 2 Associations with long COVID symptoms in full study cohort.

( a ) Fibrinogen levels at 6 months were compared between pooled LC cases (n = 295) and Recovered (n = 233) and between the Cognitive group (n = 41) and Recovered (n = 233). Box plot centre line represent the Median and boundaries of the box represent interquartile range (IQR), the whisker length represents 1.5xIQR, any outliers beyond the whisker range are shown as individual dots. Median differences were compared using two-sided Wilcoxon signed-rank test *= p  < 0·05, **= p  < 0·01, ***= p  < 0·001, ****= p  < 0·0001. Unadjusted p-values are reported. b ) Distribution of time from COVID-19 hospitalisation at sample collection applying CDC and NICE definitions of LC (n = 719) ( c ) Upset plot of symptom groups. Horizontal coloured bars represent the number of patients in each symptom group: Cardiorespiratory (Cardio_Resp), Fatigue, Cognitive, Gastrointestinal (GI) and Anxiety/Depression (Anx_Dep). Vertical black bars represent the number of patients in each symptom combination group. To prevent patient identification, where less than 5 patients belong to a combination group, this has been represented as ‘<5’. The Recovered group (n = 250) were used as controls. Forest plots show Olink protein concentrations (NPX) associated with ( d ) Cardio_Resp (n = 398), ( e ) Fatigue (n = 342), ( f ) Anx_Dep (n = 219), ( g ) GI (n = 134), and ( h ) Cognitive (n = 65). Error bars represent the median accuracy of the model.

Extended Data Fig. 3 Validation of olink measurements using conventional assays in plasma.

Olink measured protein (NPX) were compared to chemiluminescence assays (ECL or ELISA, log2[pg/mL]) to validate our findings, where contemporaneously collected plasma samples were available (n = 58). Results from key mediators associated with LC groups were validated: CSF3, IL1R2, IL2, IL3RA, TNFa, TFF2. R = spearman rank correlation coefficient and shaded areas indicated the 95% confidence interval. Samples that fell below the lower limit of detection for a given assay were excluded and the ‘n’ value on each panel indicates the number of samples above this limit.

Extended Data Fig. 4 Univariate analysis of proteins associated with each symptom.

Olink measured plasma protein levels (NPX) compared between LC groups (Cardio_Resp, n = 398, Fatigue n = 384, Anxiety/Depression, n = 202, GI, n = 132 and Cognitive, n = 60) and Recovered (n = 233). Proteins identified by PLR were compared between groups. Median differences were compared using two-sided Wilcoxon signed-rank test. * = p < 0·05, ** = p < 0·01, *** = p < 0·001, ****= p < 0·0001 after FDR adjustment. Box plot centre line represent the Median and boundaries of the box represent interquartile range (IQR), the whisker length represents 1.5xIQR, any outliers beyond the whisker range are shown as individual dots.

Extended Data Fig. 5 Unadjusted Penalised Logistic Regression.

Olink measured proteins (NPX) and their association with Cardio_Resp (n = 398), Fatigue (n = 342), Anx_Dep (n = 219), GI (n = 134), and Cognitive (n = 65). Forest plots show odds of each LC outcome vs Recovered (n = 233), using PLR without adjusting for clinical co-variates. Error bars represent the median accuracy of the model.

Extended Data Fig. 6 Partial Least Squares analysis.

Olink measured proteins (NPX) and their association with Cardio_Resp (n = 398), Fatigue (n = 342), Anx_Dep (n = 219), GI (n = 134), and Cognitive (n = 65) groups. Forest plots show odds of LC outcome vs Recovered (n = 233), using PLS analysis. Error bars represent the standard error of the coefficient estimate.

Extended Data Fig. 7 Network analysis centrality.

Each graph shows the centrality score for each Olink measured protein (NPX) found to have significant associations with other proteins that were elevated in the Cardio_Resp (n = 398), Fatigue (n = 342), Anx_Dep (n = 219), GI (n = 134), and Cognitive (n = 65) groups relative to Recovered (n = 233).

Extended Data Fig. 8 Inflammation in men and women with long COVID.

Olink measured plasma protein levels (NPX) between men and women with symptoms, divided by age (<50 or >=50years): (a) shows IL1R2 and MATN2 in the Anxiety/Depression group (<50 n = 55, >=50 n = 133), (b) shows CTSO and NFASC in the Cognitive group (<50 n = 11, >=50 n = 50). Median values were compared between men and women using two-sided Wilcoxon signed-rank test. Box plot centre line represent the Median and boundaries represent interquartile range (IQR), the whisker length represents 1.5xIQR.

Extended Data Fig. 9 Inflammation in the upper respiratory tract.

Nasal cytokines measured by immunoassay in the CardioResp Group (n = 29) and Recovered (n = 31): ( a ) shows IL1a, IL1b, IL-6, APO-2, TGFa, TFF2. Median differences were compared using two-sided Wilcoxon signed-rank test. Box plot centre line represents the Median and boundaries of the box represent interquartile range (IQR), the whisker length represent 1.5xIQR. ( b ) Shows cytokines measured by immunoassay in paired plasma and nasal (n = 70). Correlations between IL1a, IL1b, IL-6, APO-2, TGFa and TFF2 in nasal and plasma samples were compared using Spearman’s rank correlation coefficient ( R ). Shaded areas indicated the 95% confidence interval of R.

Extended Data Fig. 10 Graphical abstract.

Summary of interpretation of key findings from Olink measured proteins and their association with CardioResp (n = 398), Fatigue (n = 342), Anx/Dep (n = 219), GI (n = 134), and Cognitive (n = 65) groups relative to Recovered (n = 233).

Supplementary information

Supplementary information.

Supplementary Methods, Statistics and reproducibility statement, Supplementary Results, Supplementary Tables 1–7, Extended data figure legends, Appendix 1 (Supplementary Table 8), Appendix 2 (PHOSP-COVID author list) and Appendix 3 (ISARIC4C author list).

Reporting Summary

Rights and permissions.

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/ .

Reprints and permissions

About this article

Cite this article.

Liew, F., Efstathiou, C., Fontanella, S. et al. Large-scale phenotyping of patients with long COVID post-hospitalization reveals mechanistic subtypes of disease. Nat Immunol 25 , 607–621 (2024). https://doi.org/10.1038/s41590-024-01778-0

Download citation

Received : 11 August 2023

Accepted : 06 February 2024

Published : 08 April 2024

Issue Date : April 2024

DOI : https://doi.org/10.1038/s41590-024-01778-0

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

This article is cited by

Immune dysregulation in long covid.

• Laura Ceglarek
• Onur Boyman

Nature Immunology (2024)

Quick links

• Explore articles by subject
• Guide to authors
• Editorial policies

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.