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Manuscript Preparation

• Article Structure
• Highlight and Abstract
• Keywords and Abbreviations
• Materials and methods
• Tables and Boxes
• Supplementary Data
• Author Contribution
• Conflict of Interest
• Funding Statement
• Data Availability

Return to Information for Authors main page JXB Data Policy

All revised papers should follow the formatting guidelines below.

1. Article Structure

The following sections list the structure that the various article types published in JXB should follow. All revised papers should follow these structures.

Research Papers

• Introduction
• Discussion (or Results and discussion)
• Supplementary data (a brief, single-sentence description for each item, not the actual figures/tables or full legends)
• Acknowledgements
• Author contributions
• Conflict of interest
• Data availability
• Figure legends
• Figures, including figure legends (these should be uploaded as a separate file(s). Please find specific guidelines in the  Figures section .

Reviews (Including Flowering Newsletter Reviews, Darwin Reviews and Expert Views)

• Main text (please use headings to divide into sections)
• Data availability (for those reviews that include meta-analysis of collated datasets)
• Boxes (optional except for ‘key developments box’ for Expert Views)

Viewpoints and Insights

• Highlights (For Viewpoints)
• Introductory paragraph (please note Viewpoints and Insights do not have abstracts)
• Conflict of interest
• Boxes (optional)
• Figures, including figure legends (these should be uploaded as a separate file(s)). Guidelines for preparing figures can be found in the Figures section below.

2. Title Page

The title page should contain the title, names of all authors, and names and addresses of the institutions where the work was carried out, and an email address for each author. The date of submission, the number of tables and figures, and the word count (start of the introduction to the start of the acknowledgements, excluding materials and methods) should be listed. If the paper has supplementary data, specify the number of figures, tables or videos.

The title should be concise and informative for a broad scientific audience, with no more than 120 characters including spaces . For research papers, the title should usually state the novel scientific findings being presented. A title that poses a question can be effective. Remember this will be the most frequently read part of your paper. Please avoid abbreviations and include species names (well-known common names are acceptable).

Also add a shorter  running title of no more than 60 characters including spaces.

3. Highlight and Abstract

3.1 highlight.

Please provide a statement that, in fewer than 30 words, highlights the novelty of the paper for the non-expert. The highlight should contain the central findings of your work, as well as keywords and phrases, but should not simply repeat the title. For reviews, the highlight should state the primary objective of the review.

Highlights are used in table of content alerts online and via emails, as well as on the advance access page, so they are a useful way to ‘highlight’ your paper to readers.

3.2 Abstract

The abstract should be an engaging and informative 'stand-alone' text (without references) of no more than 200 words. Abstracts for research papers should detail why the research was undertaken; the approach and methodology if appropriate; the main findings; and the key conclusions, including wider implications. Repeat keywords and phrases as appropriate. Abbreviations should only be defined and used within the abstract if they are used three times or more in the abstract itself.

For reviews, abstracts should state the primary objective of the review and the reasoning for focusing on this objective, the principal findings of the review, and any conclusions that might be drawn from it, including implications for further research or practices. 

4. Keywords and Abbreviations

4.1 keywords.

Please list  6–10 keywords (in alphabetical order) after the abstract. Think of words people might use in searches. The main keywords should also appear in the title, abstract, highlight, and subheadings. Natural, consistent repetition of keywords will aid search engine optimisation so others can more easily find and cite the paper.

4.2 Abbreviations

Please limit the use of non-standard abbreviations, which can make the text harder to read. Generally, only use them for words used more than three times in the text but only where the shortened form will aid readability. Spell out the term on first mention: for example, ‘the International Rice Research Institute (IRRI) is developing rice varieties…’. If you do have such defined abbreviations, also list them , in alphabetical order, after the keywords.

There are a number of common abbreviations that may be used without definition. The same applies to standard chemical symbols.  Authors are asked to avoid using CK as an abbreviation for 'control' as this abbreviation is more commonly used for 'cytokinin' within plant sciences. 

5. Materials and methods

This section should provide a detailed and complete description of the materials and methods adopted in the study. The description should provide sufficient detail to enable the reader to both understand and replicate the experiments performed. Published articles may be cited instead of giving a full description, as long as authors verify that the cited article contains a reasonably complete description of the respective materials and/or method. Any deviations from the previously published method that could significantly change the outcome of the experiment must be detailed in full. Methods essential to understanding and replicating experiments should not be included in the supplementary data. The inclusion of previously unpublished primary experimental data is not permitted in all types of JXB reviews, and as such, review manuscripts should not contain a materials and methods section.

To facilitate the reproducibility of studies, authors submitting to JXB are encouraged to consider depositing novel protocols to a suitable public repository, such as protocols.io or Zenodo . Similarly, authors whose papers include code should consider uploading their code to a public repository such as GitHub . For authors submitting Technical Innovations and Community Resources that introduce new methods and tools, depositing protocols into an appropriate repository is a pre-submission requirement. Once authors have described the step-by-step protocol on their selected repository, a DOI will be issued. The DOI should be cited in the materials and methods section, enabling editors and reviewers to access the protocol. Authors can choose for their method to be kept private until their paper is published but access to editors and reviewers should be provided where possible, for example, by obtaining a shareable link for the protocol to be accessed before it is published. As well as being searchable and citable, protocols deposited in protocols.io are presented in a clear, formatted form, and new versions can be included as methodologies develop over time. Authors should not simply copy and paste all the text from within their materials and methods section when depositing in a repository.

Authors should divide the materials and methods into appropriate sections and should refer to the Styling Points for information on reporting numbers and dates, scientific names, measurements, chemicals, genes and proteins, and equations.

It is expected that  novel materials  used and described in new papers be made available for non-commercial research purposes upon publication. A statement concerning availability, or restrictions on availability, should be included in the data availability section. It is acknowledged that some materials such as enzymes, natural products, and antibodies require substantial effort to generate, and supplies may be limited—a judgement will be made on whether any restrictions on availability are reasonable. For  antibodies , also note that full source information must be provided (e.g. company, including sufficient address details to enable contact).

Quantitative real-time PCR experiments should conform to the MIQE guidelines. Please note that normalisation against a single reference gene is only acceptable if clear evidence is presented that confirms its invariant expression under the particular conditions described. Normalisation is usually conducted with at least two validated reference genes. Ideally, authors using qRT-PCR should upload a completed MIQE checklist as part of their supplementary data, providing all ‘essential’ information on the checklist. For further guidance see Graeber et al. (2011) . Please see the Minimum reporting standards section of this guide for full guidance.

6. Tables and Boxes

Each numbered table (e.g. Table 1, Table 2) should have a  concise, descriptive heading , with any further essential explanation added as a footnote. The heading should provide enough detail to understand the table without referring to the main text (i.e., it should be 'stand-alone'), although there is no need to redefine abbreviations not specific to the table or give full species names where this information has already been provided. Please create tables using the table tool in Microsoft Word. All tables should be cited in sequence in the main text.

Boxes are a way of expanding on information in a review paper without disrupting the flow of the main text. They could be about the general area (e.g., containing information indirectly related to the paper) or be more spe­cific (e.g., explaining a particular process in detail). Alternatively, boxes can be used to express an opinion or a particular point of view on a controversial topic within a review. Typically, boxes will include some text and an image or explanatory dia­gram and should have a descriptive one-line heading. Text in boxes is not includ­ed in the word limit.

The inclusion of a ‘ Key Developments Box ’ is an essential element of all Expert View articles. This box should highlight four to six papers from the past two years, each summarised in a couple of sentences. Please see our Expert View guidance for more details. Boxes are optional for other review papers (e.g. Darwin Reviews and Flowering Newsletter Reviews).

7.1 Figure legends

As well as being included underneath each figure, a list of figure legends should appear at the end of the manuscript file, after the tables. Each numbered legend (e.g. Fig. 1, Fig. 2) should begin with a concise, single-sentence description of the figure. It should go on to provide enough detail to understand the figure without referring to the main text. A description of any symbols should be given in full (do not include actual character symbols in the legend) unless a key is included in the figure itself. Micrographs must include a scale bar, ideally with the length provided on the image (but otherwise indicated in the legend).

Review and other papers that reproduce previously published figures or data, should clearly state this within the figure legend and appropriate permissions must be sought as required. Please consult the Permissions section of this guide for full details.

7.2 Figure preparation

JXB has clear requirements on the preparation of figures, including the use of composite images, figure manipulation, and data duplication. Please consult the Figure Policy prior to submitting your manuscript.

For revised manuscripts, figures should not be included in the (Word/rtf) manuscript file. They should instead be uploaded as a separate file or files. For photographs and composite figures, the preferred format is .tiff; for other figures, such as diagrams and graphs, .eps, .pdf, .doc or .ppt are also acceptable.

Each figure should be clearly labelled (Fig. 1, Fig. 2, etc.) and should include the full figure legend. They should all be cited  in sequence  in the main text.

Please make figures as clear as possible. Add labelling where there are multiple panels using capital letters (Fig. 1A, Fig. 1B, etc.). In general, use Helvetica Neue or similar sans serif font for lettering. The font size should be uniform; between 6 and 10 pt at expected final publication size is ideal. For micrographs, a scale bar with length indicated on the image is best. Line weights should be between 0.5 and 1.5 pt. Preferred symbols, in order, are closed circles, open circles, closed squares, open squares, closed triangles, and open triangles; these should be no smaller than 1 mm (height/diameter) at expected final publication size. Avoid using mathematical symbols. Resolution at final publication size should be no less than 300 dpi and at no less than 1080 pixel width.

The use of colour in figures is encouraged, and there is no charge. All figures provided in colour will be published online in colour. Please save colour figures with RGB and always follow  colour-blind-friendly practice . 

Authors should keep ready access to all original images, which should be high quality, unedited, uncropped, and high resolution. These may be requested during peer review.

7.3 Cover Images

Authors are encouraged to submit original images to be considered for the cover of the journal. Cover art may come directly from or be closely related to a paper in the issue. Please provide images in the same final format as a figure: i.e., ideally as a .tiff and at a minimum 300 dpi at publication size. Dimensions for cover images are as follows:  One image, landscape option : 216mm width x 161mm height Two images side-by-side, portrait options : 108mm width x 161mm height (each) Images may be uploaded as additional supplementary files at submission (initial submission or revision stage) or emailed to the editorial office at any time during the review process or shortly after acceptance. Please provide the manuscript ID number/title and a brief description of the image and the credit.

8. Supplementary Data

Authors may include supplementary data, but because such data do not receive a DOI and are not searchable, we encourage authors to consider instead uploading supplementary data to a public repository in the interests of transparency and accessibility. You can read more about the benefits of submitting data to a public repository, such as Zenodo , in the  JXB Data Policy  pdf. JXB also encourage all authors to consider uploading the raw data associated with their paper (for example, raw gel images, unedited micrographs, and numerical data used to generate graphs and tables) to a public repository such as Dryad , Zenodo or Figshare . More information can be found in the data deposition section of this guide.

Supplementary data for online-only publication may be submitted if they add valuable information that is not essential to a full understanding of the main paper or cannot practically be included in the paper. Supplementary data may contain figures, tables, datasets, protocols, and/ or videos. Methods essential for understanding the paper should be included in the main manuscript file and not as supplementary data. Please note that JXB does not permit the inclusion of supplementary data with review manuscripts. We instead encourage authors to deposit any large-scale collated datasets that were used for meta-analysis in a public repository (e.g Zenodo ).

To differentiate supplementary data from the figures and tables in the manuscript, please add an S to items in this section and refer to them in your manuscript as follows: Supplementary Fig. S2, Supplementary Table S1, etc. Add labelling where there are multiple panels in figures using capital letters (Supplementary Fig. S1A, Supplementary Fig. S1B, etc.).  Within each category (e.g. tables, figures, videos) all items of supplementary data should be cited in sequence in the main text.

Add a supplementary data section, immediately after the discussion in your main paper, giving each item a  brief, single-sentence description . For example:

Supplementary data

The following supplementary data are available at JXB online. Table S1 . Cowpea germplasm lines used in this study. Table S2 . Primer sequences for the amplification of DNA fragments flanking 20 randomly selected SNPs. Fig. S1 . Principal component analysis of the cowpea germplasm used in this study, showing the distribution of the eight accessions chosen for the SNP discovery panel. Fig. 2 . Osmotic stress tolerance phenotypes of independent UP12_8740-OE lines, empty vector-transformed CK lines, and wild-type plants under 6% or 10% PEG treatment. Dataset S1 . Bait library sequence used in Capture-Seq.

Ideally provide all supplementary data as a single PDF file , but complex tables and datasets are preferred as Excel files, and video files as .mov, .mpg, .avi and animated .gif files. Ensure video can be easily viewed with widely available software (e.g. Windows Media Player or QuickTime Player). For easy viewing and downloading, keep the size of the file(s) small where possible, i.e. <10 MB.

Please label each supplementary data file according to its contents, for example:

Supplementary Dataset S1; Supplementary Figure S1–S5; files with various content: Supplementary Figures S1–S2 and Tables S1–S3.

Supplementary data must be fully understandable on their own so full legends must be included in the file(s). Please do not add line numbers and do not include supplementary data in the main manuscript file. These files should be uploaded in the dedicated supplementary files upload area of the submission site.

Although these data will be subject to full peer review, they are not professionally copyedited so it is essential that authors check them meticulously.

9. Author Contribution

All research papers must include an author contribution statement, clearly identifying the contribution made by each author. JXB uses the Contributor Roles Taxonomy ( CRediT ) for describing individual author contributions to papers. All authors should agree to this statement prior to submission. The author contribution statement should appear after the acknowledgements and before the conflict of interest statement. Please see the Authorship  section of this guide for further guidance. An author contribution statement is not required for review papers.

10. Conflict of Interest

Authors are required to reveal any financial interests or connections, direct or indirect, or other situations that might raise the question of bias in the work reported or the conclusions, implications or opinions stated. This may include pertinent commercial or other sources of funding for the individual author(s) or for the associated department(s) or organisation(s), personal relationships, or direct academic competition. Existence of a conflict of interest does not preclude publication in the journal. It is the responsibility of the corresponding author to disclose on submission all pertinent commercial and other relationships of all authors. Any conflict of interest should be declared in a ‘Conflict of interest’ statement to be included in the paper after the acknowledgements and author contribution section. The statement is required for all papers. When considering whether to declare a conflicting interest or connection, please consider the conflict of interest test: is there any arrangement that would embarrass you or any of your co-authors if it were to become public after publication and you had not previously declared it? In other words, authors should declare perceived conflicts as well as direct conflicts. A detailed definition of conflict of interests can be found on the Oxford University Press Conflict of Interest page . If authors have no conflict of interest, this should be stated as follows in the conflict section: ‘No conflict of interest declared’.

11. Funding Statement

Authors must name their funding sources, or state if there are none, in a separate section entitled ‘Funding’. This should appear after the Conflict of interest section and is required for all papers. Funding statements should follow the format described below, unless specified otherwise by a funder:

• The sentence should begin: ‘This work was supported by …’
• The full official funding agency name should be given, e.g., ‘the National Cancer Institute at the National Institutes of Health’ or simply, ‘National Institutes of Health,’ not ‘NCI’ (one of the 27 subinstitutions) or ‘NCI at NIH’ (See full RIN-approved list of UK funding agencies.)
• Grant numbers should be complete and accurate and provided in brackets as follows: ‘[grant number ABX CDXXXXXX]’
• Multiple grant numbers should be separated by a comma as follows: ‘[grant numbers ABX CDXXXXXX, EFX GHXXXXXX]’
• Agencies should be separated by a semi-colon (plus ‘and’ before the last funding agency)
• Where individuals need to be specified as the recipient of certain sources of funding, the following text should be added after the relevant agency or grant number ‘to [author initials]’.
• An example is given here: ‘This work was supported by the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme [grant agreement XXXXXX] and the National Science Foundation (NSF) [grant XXXXXX to M.P.]

Authors without funding should use the following statement: ‘This research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors.’ Please see the Funding and Crossref Funding Data Registry  section of this guide for further information. 

12. Data Availability

To encourage authors to make their primary data available and facilitate the discoverability of such data, all papers submitted to JXB must have a ‘Data availability’ statement. Primary data are data that have been gathered directly from specific sources (usually experimentation or observation). The primary data are usually used to prepare the figures and tables in a paper. For papers published in JXB, deposition of primary data in a publicly available data repository is mandatory for some types of data and strongly encouraged for others. The ‘Data availability’ statement should clearly identify the locations where all types of deposited primary data can be accessed.

Mandatory deposition of primary data is required for sequence data (nucleic acids or proteins or), omics datasets (e.g. RNA-Seq, array data, proteomics mass spectra), molecular interaction data, and any other primary data where there is a field-specific, publicly available database. Deposition of these types of data is required prior to publication. More information about mandatory data deposition requirements and recommended repositories can be found in our Data Deposition Guidelines section.

While JXB does not mandate the deposition of other types of primary data, we strongly encourage authors to submit other primary data associated with their paper, such as raw phenotypic data used for statistical analysis, to a public repository (such as Dryad , Zenodo or Figshare ).

Providing large datasets in supplementary information is strongly discouraged, with authors encouraged instead to make such data available in repositories. The benefits of uploading data to repositories rather than as supplementary material can be found in the JXB Data Policy for Supplementary Data . Extra results presented in the supplementary material that do not feature in the paper must not be referenced within the data availability statement.

The ‘Data availability’ statement should report where the primary data generated within the paper are available, outline the terms of use, and provide accession codes or other unique identifiers where data have been made publicly available. It is expected that novel materials used and described in the paper are made available for non-commercial research purposes and a statement concerning availability, or restrictions on availability, should also be included in the ‘Data availability’ section. The statement should also include details of where to access previously unreported code used to generate results reported within the paper that have been made publicly available in appropriate repositories (e.g. GitHub). Authors should not use the data availability statement to refer to data that were not generated within the paper, with the exception of large-scale collated datasets subject to meta-analysis.

If data are subject to an embargo or other restrictions on availability, such as third-party restrictions, the reason for this must be provided and a release date given. As depositing primary data in a repository is encouraged we recommend authors avoid using such statements unless absolutely necessary.

The ‘Data availability' statement should be included in the manuscript after the funding statement.

Review articles may not include unpublished experimental data and the data availability statement for these papers should confirm this (e.g. "This review contains no new experimental data"). Reviews may include meta-analysis of previously published data, the sources of which should be reported in the ‘Data availability’ statement.

Examples of ‘Data availability’ statements can be found in the table below. Authors may need to combine statements to best describe the availability of primary data (for example, for primary RNA-Seq datasets uploaded to the Gene Expression Omnibus (GEO) and for general phenotypic data uploaded to Dryad).

If you have any concerns or questions about sharing your data, please contact the editorial office at  [email protected] .

More information on Data Availability Statements can be can be found on the OUP Research Data page .

13. References

Previously published work must be acknowledged by appropriate citation in the main text and a full reference list. References in research and review papers should be balanced and appropriate for setting context and demonstrating novelty. Restrictions on the number of references apply only to eXtra Botany papers: Insights and Viewpoints should have a maximum of 20 references; Editorials up to 20 references in addition to all the papers included in the issue. Please see our guides for Insights and Viewpoints  and Editorials for more details.

Attention to detail when referencing ensures correct crosslinking, enabling links from the reference to the full-text document.

In-text citation style examples:

Chen and Zhu (2015) have shown ...  ... towards the root tip (Zhu, 2014; Chen and Zhu, 2015). Note the use of chronological rather than alphabetical order. When papers are by more than two authors use et al . (e.g. Zhao  et al  ., 2015). If several papers by the same author in the same year are cited, use letters to distinguish between them (e.g. 2016  a, b ). If several papers by different authors with the same surname and in the same year are cited, use author initials to distinguish between them (Z. Zhang et al., 2017; X.Y. Zhang et al., 2017). For papers in preprint servers add Preprint (Castel et al ., 2019, Preprint)

We encourage authors to deposit all supporting data either in public repositories or in supplementary data, and hence ‘data not shown’ and ‘unpublished results’ are not permitted.

Reference List

References should be listed in alphabetical order (without numbering). As with citations, references must be accurate and follow journal style (see examples below). Note use of full journal titles (e.g., Journal of Experimental Botany not J.Exp.Bot).

For a paper with up to ten authors, list them all; for more than ten authors, list the first three followed by et al . Citation of papers from e-journals or from papers available ahead of print should include the DOI or URL rather than volume/page numbers.

Only papers published (including preprints) or in press should be cited. (If in press , a proof will need to be submitted with the paper.) Papers submitted to journals and those in preparation should not be cited in the reference list.

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Most professors and researchers prefer articles that are "scholarly" in nature.  Characteristics of scholarly journals are that they have the following qualities:

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• A list of references or sources is provided at the end of each article
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A Comprehensive Review on Cannabis sativa Ethnobotany, Phytochemistry, Molecular Docking and Biological Activities

Sohaib hourfane.

1 Research Team on Natural Products Chemistry and Smart Technology (NPC-ST), Polydisciplinary Faculty of Larache, Route de Rabat, Abdelmalek Essaadi University, Tetouan 92000, Morocco

Hicham Mechqoq

Abdellah yassine bekkali, joão miguel rocha.

2 LEPABE—Laboratory for Process Engineering, Environment, Biotechnology and Energy, Faculty of Engineering, University of Porto, Rua Dr. Roberto Frias, s/n, 4200-465 Porto, Portugal

3 ALiCE—Associate Laboratory in Chemical Engineering, Faculty of Engineering, University of Porto, Rua Dr. Roberto Frias, s/n, 4200-465 Porto, Portugal

Noureddine El Aouad

Associated data.

Data are contained within the article.

For more than a century, Cannabis was considered a narcotic and has been banned by lawmakers all over the world. In recent years, interest in this plant has increased due to its therapeutic potential, in addition to a very interesting chemical composition, characterized by the presence of an atypical family of molecules known as phytocannabinoids. With this emerging interest, it is very important to take stock of what research has been conducted so far on the chemistry and biology of Cannabis sativa . The aim of this review is to describe the traditional uses, chemical composition and biological activities of different parts of this plant, as well as the molecular docking studies. Information was collected from electronic databases, namely SciFinder, ScienceDirect, PubMed and Web of Science. Cannabis is mainly popular for its recreational use, but it is also traditionally used as remedy for the treatment of several diseases, including diabetes, digestive, circulatory, genital, nervous, urinary, skin and respiratory diseases. These biological proprieties are mainly due to the presence of bioactive metabolites represented by more than 550 different molecules. Molecular docking simulations proved the presence of affinities between Cannabis compounds and several enzymes responsible for anti-inflammatory, antidiabetic, antiepileptic and anticancer activities. Several biological activities have been evaluated on the metabolites of Cannabis sativa , and these works have shown the presence of antioxidant, antibacterial, anticoagulant, antifungal, anti-aflatoxigenic, insecticidal, anti-inflammatory, anticancer, neuroprotective and dermocosmetic activities. This paper presents the up-to-date reported investigations and opens many reflections and further research perspectives.

1. Introduction

Cannabis sativa L. is an herbaceous plant belonging to the Cannabaceae family. This plant species has many vernacular names and is known by many people as marijuana and hemp. Despite being native to Central Asia, this plant’s capacity of adaption to different climates lead to its spread all over the world [ 1 ]. The Cannabis genus is composed of a single specie named “ sativa ”, which regroup several subspecies or varieties including Cannabis sativa ssp. sativa , Cannabis sativa ssp. indica , Cannabis sativa ssp. ruderalis and Cannabis sativa ssp. afghanica . However, there is still controversy among the scientific community about the sub-classification of Cannabis species and varieties [ 2 , 3 , 4 ]. Cannabis sativa L. is one of the plants that have been used by humankind since antiquity, and many historians reported the different uses of this plant around the world [ 5 , 6 , 7 , 8 ]. The historical records show that this plant has been used as a source of fiber, food, oil, as well as for recreational and religious purposes. Additionally, several other uses have been developed through the centuries, such as livestock feed, skin and hair care [ 9 ]. Furthermore, many ethnobotanical surveys highlighted the therapeutic use of Cannabis sativa L. for the treatment of chronic pain, depression and inflammation. These activities have been justified by the original chemical composition, viz. Cannabis contains a large number of bioactive compounds with an estimation of more than 550 molecules [ 10 ]. Those compounds belong to the cannabinoid, terpenoid, stilbenoid, lignanamide, carotenoid, flavonoid and alkaloid classes [ 11 ]. The most notable compound of Cannabis remains the cannabinoids [ 12 , 13 ], a class of terpenolic compounds mainly found in the trichome cavity of female flowers [ 14 , 15 ]. Nowadays, Cannabis sativa L. is experiencing a renewed interest in many research fields, including microbiology and oncology [ 12 , 13 ]. In fact, the chemical diversity of cannabinoids proved to be very useful for targeting microorganisms such as bacteria, fungi, viruses, as well as cell components such as proteins and genes [ 16 ]. Additionally, their natural origin and low toxicity make them perfect candidates to treat hard-to-treat diseases, by solving therapeutic problems such as the resistance to antibiotics and, in the case of cancer treatment, the toxicity induced by ingestion and metabolization [ 17 , 18 , 19 ].

This review aims to present Cannabis sativa subspecies classification, description of the plant aspect and botany as well as a brief history and geographic distribution. This manuscript aims also to report and discuss the traditional uses as to both the preparation and administration modes of every part of the plant. It also aims to take into account the chemical composition of each part with the classification of the identified metabolites and their quantification, and the biological activities of Cannabis extracts and purified compounds. Finally, this review also includes the molecular docking studies of secondary metabolites previously identified in different parts of Cannabis sativa L.

2. Generalities about Cannabis sativa L.

2.1. plant nomenclature and synonyms.

Carolus Linnæus, also known as Carl von Linné (1707–1778), was the first person to frame principles for classification of living organisms into classes and sub-classes. His aim was to create a uniform international system for the identification of any living organism according to its morphological features. In this system, every organism is identified by his genera and specie names known as “binomial nomenclature”. In 1753, Carl von Linné mentioned the word Cannabis for the first time. This word comes from the Latin canna that means “reed” and bis that means “twice”, which means literally “reed with two sexes” [ 20 ]. Prior to the Linnæus nomenclature, Cannabis was widely used by different civilizations that gave it different names known as vernacular names [ 21 , 22 , 23 ]. At present, there are many local or vernacular names and various synonyms to name Cannabis. It is also known as hashish, marijuana, weed, Acapulco gold, ace, bat, bhang, log, hemp, Indian hemp, Colombian, doobie, dope (Cannabis), ganja, hydro, Jamaican, jive (sticks), joint, Maui wowie, Mexican, Panama gold, Panama red, pot, firecracker, ragweed, reefer, sativa, sinsemilla of California, spliff, Thai stick, etc. Those names and designations stay different depending on the region, country and culture. Cannabis sativa belongs to the Cannabaceae family, which includes 12 genera and 102 species, and with some species of economic importance, such as Humulus lupulus L. and Pteroceltis tatarinowii [ 24 ]. There are conflicting botanical classifications of Cannabis sativa , and the taxonomic classification of this plant has been the subject of divergences and debates. It is commonly accepted and recommended that Cannabis sativa is a single species [ 25 ], with four subspecies, namely indica , ruderalis, sativa and afghanica [ 2 , 3 , 4 , 26 ]. However, the classification criteria used for the differentiation of Cannabis sativa subspecies are often not very clear, since the chemical and morphological characteristics appear to vary according to the plant environment and pedology. In a study reported by Pacifico, et al. [ 27 ], the authors showed that the tetrahydrocannabinol (THC) content of a Cannabis sativa single species depends on the growing climate of the plant. In most cases, it is recommended to apply the name Cannabis sativa to all Cannabis plants encountered, since they all belong to the same species, and there is no agreement on the plant taxonomy [ 25 ].

2.2. Description and Botanical Aspect

Cannabis sativa L. is an annual, usually dioecious plant belonging to the Cannabaceae family [ 28 ]. It is now considered as the only species of the botanical genus Cannabis but divided into several phenotypes that can be described as subspecies or varieties [ 29 ]. Cannabis sativa has the particularity of being a fast-growing plant with a fluted stem that can reach 1 to 4 m with a diameter ranging between 1 and 3 cm ( Figure 1 a) [ 30 ]. The variation of height and diameter depends on the sub-species, environment, soil and climatic conditions [ 31 , 32 ]. The seeds are smooth, greyish ovoid or spherical in shape, 2.5 to 3.5 mm long and 2.5 to 3 mm in diameter ( Figure 1 c). Each seed contains two cotyledons rich in reserves (protein and oil), with an albumen considered particularly small compared to other plant species [ 33 ].

Cannabis sativa L. General aspect ( a ); inflorescence ( b ); seed ( c ); leaf ( d ); stem ( e ).

This plant is also characterized by long, fine flowers ( Figure 1 b). It has glandular hairs that make it fragrant and sticky [ 34 , 35 ]. At post-germination, young male and female plants cannot be distinguished. It is only during the last phase of growth, when flowers start appearing, that sex determination becomes possible [ 24 , 36 ]. The female flowers have no petals and consist of two long white, yellow or pink stigmas. Their calyx (less than 3–6 mm) envelops the ovary containing a single ovule. The female flowers appear in pairs in the axils of small leaves named bracts, these bracts contain numerous glandular trichomes where cannabinoids, mainly THC, accumulate [ 34 , 37 , 38 ]. On the other hand, the male flowers have five sepals of approximately 5 mm length, with yellow, white or green color [ 33 , 39 ]. The male plants develop small pollen sacs that serve to fertilize the female plants with hairy, resinous stigmas [ 34 , 36 , 40 ]. The Cannabis leaves are stipulate and opposite, with palmate (five to seven unequal), elongated and spiny segments with toothed margins ( Figure 1 d). Towards the top of the axis, the leaves become alternate and are inserted on the stem in an opposite arrangement every 10–30 cm [ 39 ]. These plants have cystolithic, tectorial and resin-secreting hairs; the latter have a voluminous base ending in a cluster of several cells, with each one secreting resin [ 39 ]. The root is taproot with a length of up to 30 cm, but the lateral roots reach 20 to 100 cm. In addition, in peaty soils, the lateral roots are more strongly developed, and the main root grows to a depth of 10–20 cm [ 41 ]. The growth rate of the root system is quite slow in the initial stages of vegetation, in contrast to the aerial part of the Cannabis plant, which grows intensively and rapidly [ 41 ].

2.3. Geographic Distribution and History

In nature, Cannabis is an annual flowering plant. This means that it completes its life cycle, from germination to seed production, in one year [ 42 ]. Cannabis can grow in a vast majority of climates ( Figure 2 ). From its region of origin, it appreciates calcareous and nitrogenous soils with a neutral or slightly acidic pH [ 43 , 44 ].

Geographic distribution of Cannabis sativa L.

This species originates from equatorial and subtropical regions, mainly from central Asia [ 1 ], where two places seem to be its cradle: the foothills of the Himalayas and the plains of the Pamir (a high mountain range centered in eastern Tajikistan with extensions into Afghanistan, the Republic of China and Kyrgyzstan) [ 45 ]. However, this plant has a wide geographical distribution growing up in Canada, United States of America, Europe and Africa. Cannabis is an ancient plant but the craze it has generated over (at least) the last century has greatly changed its face and even the face of the world. It is probably the first plant domesticated by humankind [ 46 ]. Many historical reports prove that this plant had been cultivated worldwide for thousands of years. The oldest documented evidence of Cannabis cultivation is a 26,900 B.C. hemp rope found in the Czech Republic [ 47 ]. Some of the earliest known prolific uses of Cannabis began in China around 10,000 B.C., where Cannabis was used to make clothing, rope and paper [ 48 ]. Further traces were reportedly found at the Neolithic site of Xianrendong on Chinese ceramics dating back to 8000 B.C. and decorated with hemp braided fibers. Between 8000 and 300 B.C., Cannabis was also cultivated in Japan and employed to make cloth fiber and paper [ 49 , 50 ]. However, the earliest reference of Cannabis psychotropic use goes back to 2700 B.C. It has been mentioned in the Chinese pharmacopoeia of the Emperor Chen Nong, where it is recommended as a sedative and remedy for insanity. Cannabis was also mentioned on the Ebers Papyrus of pharaonic Egypt back to 1550 B.C. as remedy for vaginal inflammations [ 51 ]. Yet, it was mentioned in Greek medicine, in the writings of Dioscorides, who underlines the psychotropic properties of the plant and already Galen fears that “it hurts the brain when we take too much” [ 52 , 53 ]. In India, it was one of the five magical plants used in religious rituals in the form of fumigation. In fact, around 1300 B.C., the stimulating and euphoric powers of bhanga (hemp in Sanskrit) were praised by the Indo-Aryans in one of the four holy books, the Atharva Veda [ 54 ]. Back to the European Continent, and around 700 B.C. in Marseille (France), Cannabis was used for rope manufacturing. The name Cannebiere (important avenue of the city) testifies of the importance of Cannabis at that time [ 35 ]. Jamestown settlers introduced Cannabis to colonial America in the early 1600s for the manufacture of rope, paper and other fiber products. This plant was so important that American presidents George Washington and Thomas Jefferson grew Cannabis [ 38 ]. The question of when and how Cannabis originated in the new world is still very controversial indeed. Cannabis was discovered in native American civilizations prior to Columbus’ arrival [ 55 ]. William Henry Holmes’ 1896 report “prehistoric textile art of the Eastern United States” indicated that Cannabis originated with native American tribes of the Great Lakes and Mississippi valley [ 56 ]. Cannabis products from pre-Columbian indigenous civilizations have also been found in Virginia [ 57 ]. Cannabis was an important crop in the United States until 1937, when the Marihuana Tax Act all but wiped out the American hemp industry. During World War II, Cannabis experienced a resurgence in the United States of America, as it was widely used to manufacture military items ranging from uniforms to canvas and rope [ 57 ]. At present, the most notable development in Cannabis production around the world is the rise of indoor cultivation, particularly in Europe, Australia and North America. This type of cultivation gives rise to a very lucrative trade, which is increasingly a source of profit for local organized crime groups [ 58 ].

3. Methodology

Relevant information about Cannabis sativa L. was collected from various scientific sources including SciFinder, ScienceDirect, PubMed and Web of Science. The targeted databases were probed with “Cannabis sativa”, “botany”, “history”, “ethnobotany”, “traditional use”, “phytochemistry”, “pharmacology”, “bioactivity”, “bioinformatic” and “in-silico prediction” as keywords. Thus, available articles were collected, summarized in tables and analyzed. In addition to that, we report up-to-date studies of Cannabis sativa ethnopharmacology, chemical composition, pharmacology and molecular docking simulations; this review aims to give a subjective critique to reported articles and offer perspectives for further investigations on Cannabis phytochemistry and pharmacology.

4. Results and Discussion

4.1. traditional uses of cannabis sativa l..

Cannabis sativa L. has been used in a wide variety of fields and showed a high usability potential with many applications including manufacturing of tools, construction, cosmetics, medication, shelter insulation, papermaking, human nutrition, animal feed, agrofuels, composite materials in association with plastics, etc. [ 5 , 6 , 7 , 8 ]. Table 1 summarizes the parts of the plant and their traditional uses.

Traditional uses of different parts of Cannabis sativa .

The analysis of collected data shows that seeds and leaves are the most used parts for medication. Many studies reported the use of Cannabis seed as food. It is used for the production of pasta, gluten-free flour characterized by a nutty taste, beer and oil [ 72 , 73 ]. In addition to their use for human nutrition [ 59 ], the seeds are also used to treat nausea, vomiting, stimulate the appetite of AIDS patients, cancer and hepatitis C. It is also applied as a muscle relaxant, for weight control, lung capacity enhancer and as an analgesic, anxiolytic, antiepileptic, antiemetic and against neurological pain [ 60 ]. These seeds have also a cosmetic use, mainly for hair fortification as a hair serum by external application of seed powder [ 61 ]. Beyond their potential value as medicine or food, Cannabis sativa seeds have recently been used to treat contaminated groundwater, since hemp seed protein powder proved to be more effective than other plant protein sources for chelating perfluoroalkyl and polyfluoroalkyl substances, known as “eternal chemicals” [ 74 ]. Those Cannabis proteins proved to be very useful for the treatment of salt-contaminated soils as well [ 75 ].

The leaves are externally used as poultice to treat eczema [ 61 ] and subcutaneous tissue disorders [ 62 ]. They are also orally consumed by local people for the treatment of central nervous system (CNS) disorders such as schizophrenia, gout, arthritic pain, bloating, coughing and mucus [ 63 , 64 ]. Cannabis leaves are among the most iconic symbols of modern stoner culture; their shape is frequently associated with the recreative use of this plant. Several studies described the traditional uses of Cannabis leaves, and those applications include the treatment of a wide range of health problems such as hypertension, rheumatoid arthritis, itching, cancer, snake and scorpion poisoning [ 66 ], as well as gastric and circulatory system disorders [ 62 ]. These leaves have also been described as strong analgesics, sedatives and narcotics [ 76 ]. Other studies described the use of Cannabis sativa stem fibers as firewood [ 62 ], for construction, tools, clothes, paper and rope manufacturing [ 5 , 59 ]. These fibers are obtained by a process called defibration, which can be briefly described as a stem beating and grinding. During this process, two co-products are obtained, namely chenevotte and Cannabis dust. It is important to highlight that the quality of the fiber decreases with the maturity of the plant, since the fibers become harder and coarser. In addition to the aerial parts of Cannabis plant, the root parts are also used for medication. They are used in particular for the treatment of joint pain, skin burns, inflammation, vermin and erysipelas infection [ 70 ]. These roots are also orally used in the form of juice to relieve issues stemming from childbirth, postpartum and hemorrhage [ 70 ]. Among the most cited uses of Cannabis sativa , the psychoactive remains the most present. The leaves and inflorescences have been consumed as a narcotic in different forms and have been prepared using different methods; for instance, the leaves are smoked or prepared, the inflorescences or resin are processed into charas or attar, hashish, ganja and plant powder, whereas leaves, inflorescences and shoots are used to prepare drinks (e.g., bhang, thandai, tandai, etc.) [ 5 , 77 ]. Moreover, Klauke, et al. [ 62 ] reported the religious use of Cannabis drinks. This plant preparation, referred to as traditional bhang drink, is highly consumed during Indian festivals such as Shivaratri and Holi [ 78 ]. Some ethnobotanical surveys described the used of the whole Cannabis plant. The aerial parts are mostly used for the treatment of mental disorders and nervous-system-related conditions. However, the most common use of those parts is for the treatment of gastric disorders, diabetes, scarring and asthma [ 71 ]. Conversely, some studies reported the appetite-stimulating, antidysentery and antidiarrhea effects of Cannabis inflorescences, but omitted the mention of preparation and administration modes [ 62 ]. The analysis of ethnobotanical findings shows that Cannabis sativa is a plant that was integrally exploited by local populations; some traditional uses are common to many countries whereas others are specific to some cultures. One can cite the psychotropic, medicinal and cosmetic purposes found all over the world, in contrast with the religious uses exclusively reported in Asian and Latin American countries. The previously reported activities and proprieties are mainly due to the presence of metabolites with interesting chemical structures; the ethnobotanical uses of Cannabis attracted phytochemists to investigate its chemical composition. The first compound to be identified and isolated from Cannabis was cannabinol at the end of the 19th century [ 79 ].

4.2. Chemical Composition of Cannabis sativa L.

Numerous studies have shown the importance of Cannabis secondary metabolites as well as their roles. This plant offers a rich reservoir of bioactive molecules that can be used for the production of pharmaceutical, nutraceutical and cosmetic products. Table 2 regroups the chemical composition of Cannabis sativa seeds, flowers, leaves and resin.

Chemical composition of Cannabis sativa different plant parts.

Cannabidiol (CBD); cannabidivarine (CBDV); cannabicitran (CBTC); cannabigerol (CBG); cannabichromene (CBC); cannabinol (CBN); δ9-tetrahydrocannabinol (δ 9 -THC); cannabidiolic acid (CBDA); tetrahydrocannabinolic acid (THCA); delta-8-tetrahydrocannabinol (δ 8 -THC); cannabigerol (CBG); cannabigerol acid (CBGA); cannabielsoin (CBE); cannabicitran (CBTC); cannabiripsol (CBR); total cannabidiol (tCBD); total cannabigerol (tCBG); total tetrahydrocannabinol (tTHC).

The chemical investigations conducted in different Cannabis sativa plant parts shows that terpenes, polyphenols and cannabinoids are the main represented secondary metabolites. Terpenes are represented by more than 100 molecules identified in the flowers, roots and leaves, as well as in the secretory glandular hairs considered as the main production site [ 11 , 98 ]. Furthermore, more than 20 polyphenols have been identified, and they are mainly flavonoids belonging to the flavone and flavonol subclasses [ 99 ]. Concerning the cannabinoids, they are among the most represented metabolites of Cannabis despite being represented by less than 20 molecules.

Cannabis seeds contain approximately 40% oil, 30% fibers and 25% proteins [ 85 , 100 ]. Those oils are rich in triacylglycerols (TAGs) represented by 18 different molecules; the predominating tags were LLL and OLLD with respective values of 23 and 19% [ 100 ]. Moreover, those oils contain high amounts of polyunsaturated fatty acids, which represent approximately 80% total fatty acids [ 101 ]. The fatty acid composition is characterized by the predominance of linoleic acid with range values of 45–60%, followed by oleic acid and palmitic with respective range values of 15–40% and 5–6% [ 80 , 81 ].

Hemp seeds contains also considerable amounts of polyphenols and tocopherols. According to Babiker, et al. [ 81 ], the hydroalcoholic extract of Cannabis seeds contained many polyphenols such as gallic acid (12.9 ± 18.3 mg/100 g) and catechin (6.0 ± 5.2 mg/100 g), whereas Moccia, et al. [ 83 ] reported the presence of additional polyphenols, namely quercetin-o-glucoside, n-trans-caffeoyltyramine and rutin. Moreover, another sub-class of polyphenols known as cannabisins have been reported on the seed hydroalcoholic maceration by Moccia, et al. [ 83 ]. The last authors described the presence of 11 molecules, namely cannabisin A, B, C, D, E, F, G, I, M, N and O, although these compounds have not been quantified. Concerning the tocopherols, four different isomers have been identified: the lead tocopherols are γ- and δ-tocopherols with 426 and 33 mg/kg, respectively [ 82 , 84 ].

Cannabinoids are a group of C21 or C22 terpenolic compounds mainly produced in Cannabis. They have also been reported in other plant species of the genus Radula and Helichrysum [ 102 ]. These cannabinoids are mainly present in leaves and inflorescences. However, Marzorati, et al. [ 103 ] reported the presence of some cannabinoids in the hydroalcoholic maceration of seeds, and Stambouli, et al. [ 104 ] reported the presence of mainly the cannabinoids tetrahydrocannabinol (THC), cannabidiol (CBD) and cannabigerol (CBG) in seed oil. These results are probably due to contamination of seeds by inflorescence resins, since cannabinoids are not produced or transported to the seeds [ 105 ]. Moreover, phytosterols, a group of lipids with a structure similar to cholesterols, mainly found in vegetable oils, have been identified in seed oils. Aiello, et al. [ 85 ] and Stambouli, et al. [ 84 ] mentioned the presence of this class of metabolites represented by the β-sitosterol, with 65–90%, and campesterol, with 6–17%. Furthermore, Stambouli, et al. [ 84 ] described the presence of an additional phytosterol known as δ-5-avenasterol, with a value of 7.8%. The carotenoids are among the less represented compounds in Cannabis seeds. They have been identified in the seed ethanolic extract, whereas lutein and β-carotene have been reported as major sterols, with 2.5 and 0.5 mg/100 g of dry weight, respectively [ 82 ].

The composition in hemp seeds of fatty acids, polyphenols, phytosterols, proteins and fibers, and precisely the presence of insoluble fibers in addition to a wide variety of minerals represented by phosphorus, potassium, magnesium, sulfur and calcium, as well as modest amounts of iron and zinc (an important enzyme cofactor for immunity and food absorption), widely justifies its biological proprieties and importance for human nutrition [ 106 , 107 ].

The Cannabis leaves contain terpenes, polyphenols, cannabinoids and alkaloids. The leaf essential oils are characterized by the presence of (e)-caryophyllene (28.3 ± 4.1%), α-humulene (9.3 ± 1.1%), β-selinene (4.7 ± 0.9%), caryophyllene oxide (4.3 ± 0.9%), α-selinene (3.1± 0.6%) and α-trans-bergamotene (2.7 ± 0.5%) [ 86 ]. These volatile terpenes are generally found in photosynthetic plant leaves and plays the role of protection from parasites and water loss [ 108 ]. The polyphenols of Cannabis leaves are mainly flavonoids and glycosides with apigenin and luteolin. These flavonoids represent 4 mg per g of the plant material. Elsewhere, the cannabinoids of Cannabis leaves had been described by Nagy, et al. [ 86 ]. The authors reported the presence of cannabidiol (CBD), cannabidivarine (CBDV), tetrahydrocannabinol (THC) and cannabichromene (CBC) with 11, 0.8, 0.7 and 0.5%, respectively. Moreover, Zagórska-Dziok, et al. [ 87 ] reported the lead presence of CBDA and CBD, with respective values of 150 and 31 mg/g of dry matter. Similarly to the cannabinoids, the alkaloids protect plants from predators and regulate plant growth [ 109 ]. These alkaloids have been described in cannabis leaves; Fasakin, et al. [ 88 ] described the presence of some alkaloids, with cannabisativine (410.30 μg/g), cannabimine C (376.12 μg/g) and anhydrocannabisativine (218.11 μg/g) as lead compounds.

The flowers show a chemical composition qualitatively similar to the leaves. They are mainly composed of terpenes, polyphenols and cannabinoids. The Cannabis flower terpenes have been identified in the essential oils obtained by hydrodistillation. According to Nagy, et al. [ 86 ], these essential oils are mainly composed of (E)-caryophyllene, α-humulene, β-selinene and α-selinene, with values of 29, 10, 4 and 3%, respectively. Additionally, Fischedick, et al. [ 110 ] reported the presence of mono- and sesquiterpenes in Cannabis flower essential oils. This study featured the quantification of monoterpenes and proved that they dominate the chemical composition, with a concentration of 28.3 mg/g of dry weight. These monoterpenes are represented by d-limonene, β-myrcene, α- and β-pinene. Furthermore, sesquiterpenes are represented by β-caryophyllene and α- humulene. These sesquiterpenes represent a concentration ranging between 0.5 and 10.1 mg/g of dry weight [ 110 ]. Polyphenols and cannabinoids have also been identified in the methanolic extract of flowers. The main polyphenols of flowers are quercetin di-c-hexoside and luteolin c-hexoside-2″-o-hexoside with 2.55 mg/g and 1.01 mg/g, respectively [ 86 ]. On the other hand, the cannabinoids are represented by cannabidiol (CBD), tetrahydrocannabinol (THC), cannabidivarine (CBDV), cannabicitran (CBTC) and cannabichromene (CBC), with respective values of 25, 1.5, 1.5, 1 and 0.2% [ 86 ]. The cannabinoidic composition of flowers fluctuates due to several environmental factors (e.g., temperature, soil nutrients, desiccation, insect predation and ultraviolet radiation) and genetic factors (varieties and heredity) [ 111 ]. According to Yang, et al. [ 112 ], the total THC, CBD and CBG increases significantly as the flowers matures, reaching the highest concentration during 6 to 7 weeks after anthesis.

The inflorescences of Cannabis have also been evaluated for their composition, and similar classes of metabolites have been described. These compounds belong to the same classes previously reported for the leaves and flowers. The essential oils of inflorescences have been reported in two research manuscripts. The first, published by Laznik, et al. [ 90 ], reported the presence of transcaryophyllene (38.2 ± 1.7%), nerolidol (12.7 ± 1.2%) and α-pinene (11.8 ± 0.4%), whereas the second, published by Pieracci et al. [ 91 ], reported the presence of β-caryophyllene (14.4 ± 0.89%), caryophyllene oxide (7.0 ± 1.06%) and α-humulene (5.3 ± 0.10%) as lead compounds. This variation is probably due to the variation of Cannabis sativa subspecies, cultivars and/or geographic, climatic and pedologic parameters. Laznik, et al. [ 90 ] also reported the chemical composition of inflorescence methanolic extract and concluded that this extract contained mainly cannabinoids represented by cannabidiolic acid (CBDA) and cannabidiol (CBD) with 9.5 and 8.7%, respectively.

Several studies have also described the composition of other Cannabis sativa L. parts, including roots, stem and resin. Sakakibara, et al. [ 113 ] and Lesma, et al. [ 114 ] reported the presence of phenolic amides and lignanamides in the fruits and roots of Cannabis. These compounds belong to the classes of lignans and polyphenols and are mainly cannabisin (A, B, C, D, E, F and G) and grossamide [ 99 ]. Triterpenes have also been found in Cannabis roots in form of friedelin and epifriedelanol [ 115 ]. The glandular secretory hairs are the main site of resin production. The latter is a yellow and sticky substance which contains the active principles. Some studies have shown its chemical composition. Stambouli, et al. [ 97 ] reported a high concentration of THC with a value higher than 20%. More recently, Elkins, et al. [ 96 ] identified and confirmed the high content of cannabinoids including cannabidiol (CBD) with a concentration of 72.12 μg/mL, followed by tetrahydrocannabinol (THC) with 48.02 μg/mL and cannabichromene (CBC) with 4.78 μg/mL. The concentrations of Cannabis sativa secondary metabolites depend on tissue type, age, variety, growing conditions (soil nutrition, temperature, humidity, UV radiation or light), harvest time (maturity) and storage conditions [ 80 , 111 , 116 , 117 ]. The Cannabis seeds are rich in oils and starch mainly composed of terpenes [ 118 ]. The seed oils serve as food for the young plant during the early stages of germination. In fact, the plant embryo needs a source of nutrition prior to its contact with soil and air [ 119 ]. Moreover, the leaves, flowers and inflorescences are rich in volatile terpenes, polyphenols and cannabinoids. These metabolites are generally involved in defense against ultraviolet radiation or aggression by pathogens. Unlike polyphenols, cannabinoids are more interesting due to their chemical structures and representativeness of Cannabis genus. The study of chemical composition of Cannabis plant parts is very important for the determination of biological activities. It is possible to predict a biological activity from the exhaustive chemical composition of an extract using bioinformatics tools such as molecular docking. This approach has been widely applied on Cannabis secondary metabolites, as described in the next section.

4.3. Molecular Docking Studies of Cannabis sativa L.

Molecular docking is a computational approach aiming to predict potential interactions between one or more ligands and a protein [ 120 ]. This approach predicts the optimal spatial conformation and orientation of the ligand within a protein active site, in addition to the determination of the interaction mode and binding affinity represented by a score. Molecular docking is the in silico equivalent of real high-throughput screening in which many molecules are tested against biological targets. The main goal of this approach is to discriminate active and inactive agents, in order to identify new molecules that will serve as a starting point for medicinal chemists [ 121 ].

The process of molecular docking can be subdivided into two basic steps, namely docking and scoring [ 122 ]. Docking is the step in which all possible spatial interactions between a ligand and a receptor are tested in order to identify the optimal interactions, whereas the binding affinity between the ligand and the receptor are quantified in scoring, and a score is given to the poses recorded after the docking phase.

Currently, there are more than 30 available docking software packages [ 123 ]. Most of them are also designed for virtual screening (independent dockings of multiple ligands with a protein). The three most frequently cited docking tools are AutoDock, GOLD and flex; they represent 27, 15 and 11% of the references, respectively [ 124 , 125 ].

Despite being very useful for guiding the selection of bioactive molecules for in vitro testing, the molecular docking simulations remains a prediction and can sometimes give erroneous results. They can be expressed either as a false negative, when an active molecule gives low docking affinity, or a false positive, when a non-active molecule is identified as a strong ligand [ 126 ]. However, this approach remains useful as a pre-investigation predictive tool. Concerning the Cannabis plant, several molecular docking studies have been reported on the different classes of metabolites, namely cannabinoids, terpenes, polyphenols, flavonoids, lignanamides, alkaloids, vitamins and proteins. These classes have been probed against a large number of specific enzymes that instigate important roles in different physiological processes (digestion, nerve conduction, hormone synthesis, etc.). Results of molecular docking are expressed in kcal/mol, and the lowest values correspond to higher affinity between a ligand and a protein. The studies reported in the bibliography are summarized in Table 3 .

Molecular docking of Cannabis sativa L. compounds.

Cannabichromene (CBC); cannabichromenic acid (CBCA); cannabichromevarin (CBCV); cannabichromevarinic acid (CBCVA); cannabicitran (CBTC); cannabicoumaronone (CBCN); cannabicyclol (CBL); cannabidiol (CBD); cannabidiol-c4 (CBDC4); cannabidiolic acid (CBDA); cannabidiorcol (CBDC); cannabidivarin (CBDV); cannabielsoin (CBE); cannabielsoin (CBL); cannabigerol (CBG); cannabigerolic acid (CBGA); cannabigerovarin (CBGV); cannabinodiol (CBND); cannabinodivarin (CBVD); cannabinol (CBN); cannabinol methyl ether (CBNM); cannabiripsol (CBR); cannabitriol (CBT); cannabivarin (CVN); tetrahydrocannabinol (THC); tetrahydrocannabivarin (THCV); δ-8-tetrahydrocannabinol (Δ-8-THC); δ-9-tetrahydrocannabinol (Δ-9-THC); δ9-tetrahydrocannabinolic acid (Δ-9-THCA).

4.3.1. Pesticidal Activity

Cholinesterase is an enzyme that catalyzes the hydrolysis reaction of a choline ester (acetylcholine, butyrylcholine) into choline and acetic acid. Acetylcholine is a well-known excitatory neurotransmitter that causes muscle contraction and stimulates the release of certain hormones. The inhibition of choline esterase causes the disfunction of nerve impulse transmission inducing mortality, and this activity is highly coveted for the elimination of pests and insects. The molecular docking of cannabis secondary metabolites against two types of cholinesterase, namely acetylcholinesterase (ACHE) and butyrylcholinesterase (BCHE), was described by Karimi, et al. [ 127 ]. In this article, the authors tested compounds belonging to the cannabinoid, flavonoid, terpene and phytosterol classes. For acetylcholinesterase, cannabioxepane, δ-9-THCA, Δ-8-THC and CBN showed scores lower than −10 kcal/mol, whereas cannabioxepane, CBL, CBN, CBT and Δ-8-THC showed scores lower than −8.5 kcal/mol against butyrylcholinesterase. Another investigation reported by Nasreen, et al. [ 128 ] reported the acetylcholinesterase docking with some cannabinoids, and CBD showed the best score with a value of −14.38 kcal/mol. Despite using the same docking software and acetylcholinesterase three-dimensional structure, the CBD score is better than the ones previously reported by Karimi, et al. [ 127 ]. This difference of results can be explained by the variation of docking parameters, such as the gridbox dimensions, position and the exhaustiveness.

4.3.2. Antimalarial and Anti-Leishmania Activities

The antimalarial activity of cannabinoids from Cannabis sativa was reported by Quan, et al. [ 129 ]. Their studied cannabinoids were docked against plasmodium falciparum dihydrofolate reductase-thymidinesynthase to recognize the potential binding affinities of these phytochemicals. Furthermore, the in silico antileishmanial activity of phytochemicals from Cannabis sativa has been well reported in the literature. In the studies conducted by Ogungbe, et al. [ 130 ], a molecular docking analysis was performed to examine the potential leishmania protein targets of plant-derived antiprotozoal polyphenolic compounds. A total of 352 phenolic phytochemicals—including 10 aurones, 6 cannabinoids, 34 chalcones, 20 chromenes, 52 coumarins, 92 flavonoids, 41 isoflavonoids, 52 lignans, 25 quinones, 8 stilbenoids, 9 xanthones and 3 miscellaneous phenolic compounds—were used in the virtual screening study with 24 leishmania enzymes (52 different protein structures from the protein data bank). Notable target proteins were leishmania dihydroorotate dehydrogenase, n-myristoyl transferase, phosphodiesterase b1, pteridine reductase, methionyl-trna synthetase, tyrosyl-trna synthetase, uridine diphosphate-glucose pyrophosphorylase, nicotinamidase and glycerol-3-phosphate dehydrogenase. The results showed that docked polyphenols can be considered as promising drug leads deserving further investigation.

4.3.3. Antiviral Activity

Despite great advances in medical and pharmaceutical research in recent years, diseases caused by viruses have remained a huge burden on public health like coronavirus, in particular SARS-CoV-2. In silico studies, including molecular docking, have repeatedly proved to be useful in addressing the particular challenges of antiviral drug discovery. A study published by Srivastava et al. [ 131 ] showed that cannabidiol may have an good affinity with a COVID-19 protease, and such affinity is represented by a score value of −7.10 kcal/mol. In another study, cannflavin exhibited a better score against an HIV-protease with a score of −9.70 kcal/mol [ 132 ].

4.3.4. Anti-Inflammatory Activity

Inflammation is a natural body reaction to injury and infection. It is mainly due to the deployment of immune system cells to the site of the injury or infection. The four symptoms of inflammation are heat, redness, swelling and pain. However, anti-inflammatory drugs are used to combat inflammation regardless of the cause of the inflammation. They are symptomatic treatments, i.e., they do not eliminate the cause of the inflammation but only its consequence and have an analgesic action. In a recent study, a panel of proteins, including the cellular tumor antigen p53, the essential modulator of NF-KB, the tumor necrosis factor (TNF) receptor, the transcription factor p65, NF-KB p105, the NF-k-b α inhibitor, the inhibitor of nuclear factor k-b kinase α subunit and the epidermal growth factor receptor, were identified as a primary target implicated in cannabidiol (CBD) anti-inflammatory activity. This finding was supported by molecular docking, which showed interactions between the major proteins and CBD. In addition, several signaling pathways, including TCF, toll-like receptors, mitogen-activated protein kinases, nuclear factor kappa, activated b-cell light chain activator and nucleotide-binding oligomerization domain receptors, were identified as key regulators in mediating the anti-inflammatory activity of CBD [ 136 ].

4.3.5. Anticancer Activity

Several molecular docking studies were interested in the potential anticancer activity of molecules derived from Cannabis sativa L. The molecular docking performed on placental aromatase cytochrome p450 was reported by Baroi, et al. [ 138 ]. These authors reported that cannabinoids, mainly cannabidiorcol and cannabidivarin, potentially bind with the best binding energies of −9.03 kcal/mol and −8.34 kcal/mol, respectively. In another study, molecular docking calculations were also performed to investigate the binding affinity of cannabinoids in the active site of crystal structure of the DLC1 RhoGAP domain in liver cancer 1. According to the performed calculations, cannabichromene and cannabidiolic acid showed promising results with regard to binding affinity to the target GTPase-activating proteins [ 137 ]. These compounds are held within the active site by a variety of non-covalent interactions, in particular hydrogen bonds, involving important amino acids. Similarly, cannabinoids were docked to predict their anti-inflammatory and anticancer activity. The researchers reported that cannabigerol and cannabichromene potentially bind to arachidonate 5-lypoxygenase with a binding energy score of −5.34 kcal/mol and −5.14 kcal/mol, respectively [ 142 ]. Cannabis flavonoids were also docked against topoisomerase II α to investigate the binding affinity of flavonoids in the active site of topoisomerase II α. The results showed that docked flavonoids can be considered as promising drug leads, thus deserving further investigation.

4.3.6. Antiepileptic Activity

A study published by Li, et al. [ 146 ] aimed to examine the mechanism of action of Cannabis on epilepsy, focusing on key compounds, targets and pathways. The molecular docking simulations were applied to identify the active ingredients and potential targets of Cannabis in the treatment of epilepsy. Topological analysis showed that cannabinoid receptor 1, albumin and glycogen synthase kinase-3 β (cnr1, alb and gsk3b) were the key targets with intense interaction. The results showed that cannabinol methyl ether could be the lead compound on the basis of molecular docking against docked protein targets. Therefore, these studies shed holistic light on the active components of Cannabis, which contributes to the search for lead compounds and the development of new drugs for the treatment of neurological diseases.

4.3.7. Neuroprotective Activity

Neuroprotective agents target the various deleterious mechanisms that occur in cerebral ischemia, with the aim of limiting the extension of the ischemic heart. It has been shown that the cannabinoid substances contained in the Cannabis sativa plant have great potential in a wide variety of therapeutic applications. However, its neuroprotective capacity has been the most studied in diseases such as Alzheimer’s disease, Parkinson’s disease, Huntington’s disease, multiple sclerosis and amyotrophic lateral sclerosis [ 151 ]. The only example of proteins as infectious agents leading to neurodegenerative disorders was the prion protein. Since then, the characteristic self-seeding mechanism of the prion protein has also been attributed to other proteins associated with neurodegenerative diseases, notably amyloid-β (aβ). Modelling with the aβ monomer and pentamer revealed that cannabinoids interacted with the aβ protein mainly through steric interactions and hydrogen bonds. The results showed that CBG bound with the highest affinity of all the docked cannabinoids. The authors of this study reported that this was mainly due to the presence of a geranyl side chain in the CBG structure, as this side chain is associated with increased lipophilicity and may, therefore, increase the propensity to bind in the hydrophobic groove of the pentamer [ 152 ].

4.3.8. Dermocosmetic Activities

Dermocosmetic products are on the borderline between cosmetic products and medicines. Considered as cosmetic products, they are mainly used to ensure photoprotection of the skin, i.e., to limit the effects of its exposure to solar radiation, but also to improve the appearance of dry or aged skin, reduce inflammatory dermatological conditions (acne, couperose, seborrheic and atopic dermatitis, psoriasis, etc.), as well as for the care of nails and hair [ 153 ]. Furthermore, tyrosinase is a key enzyme in the process of melanogenesis (the biosynthesis of melanin). The dermocosmetic potential of Cannabis polyphenols has been reported, and the results showed a good affinity with the tyrosine phosphatase-1b with a free energy score of −24.34 kcal/mol [ 149 ]. Similarly, Cannabis alkaloids have been shown to possess affinity with tyrosinase with score a value of −3 kcal/mol [ 154 ].

4.4. Biological Activities of Cannabis sativa L.

Studies with Cannabis sativa L. have shown the presence of several biological activities, such as antioxidant, antibacterial, anticoagulant, insecticide, anticancer, anti-aflatoxigenic, antifungal, cytotoxic, anti-elastase, anti-collagenase, anti-acetylcholinesterase, anti-inflammatory, neuroprotective (anti-Alzheimer’s, anti-epilepsy and anti-Parkinson’s) and dermocosmetic (anti-tyrosinase, anti-collagenase and anti-elastase). Table 4 summarizes the biological activities of different Cannabis sativa parts according to the literature.

Biological activities of different parts of Cannabis sativa L.

IPC 50 : concentration providing 50% inhibition of lipid peroxidation/CC 50 : concentration providing 50% metal chelating activity. IC 50 : concentration of the sample that inhibits 50% expressed in mg/mL/ID: inhibition diameter (mm). EDTA: ethylenediaminetetraacetic acid./GAE: gallic acid equivalent./RE: rutin equivalent./CE: caffeic acid equivalent./TE: trolox equivalent. EDTAE: ethylenediaminetetraacetic acid equivalent./MIC: minimum inhibitory concentration./MBC: minimum bactericidal concentration. MCF-7: estrogen-dependent breast cancer cells./MDA-MB-468: triple-negative breast cancer cells./CACO-2: colorectal adenocarcinoma cells./MZ-CHA-1: cholangiocarcinoma cells.

4.4.1. Antioxidant Activity

An antioxidant is a molecule that slows down or prevents the oxidation of molecules that can play an important role in an organism metabolism. Cannabis sativa L. proved to have plenty of antioxidant substances. The antioxidant activity of this plant has been widely reported in the literature and it was determined by using many assays, including the free radical scavenging method (DPPH), oxygen radical absorbance capacity (ORAC), ferric reducing ability of plasma (FRAP) and 2,2′-azino-bis (3-ethylbenzothiazoline-6-sulphonic acid) (ABTS), as well as other methods such as phosphomolybdenum and metal chelation. The antioxidant activity of Cannabis sativa has been reported in the plant seeds, leaves and aerial parts. Research on the antioxidant effects of Cannabis sativa L. seeds have been well reported in the literature. Manosroi, et al. [ 155 ] described the antioxidant activity of ethanolic extract using different tests, namely DPPH, chelating assay and lipid peroxidation inhibition. The obtained results showed that the seed organic extract exhibited strong antioxidant capacity, suggested by IC 50 (low inhibitory concentration at 50%) values with 14.5 mg/mL for DPPH, 1.9 mg/mL for chelating assay and 92.7 mg/mL for lipid peroxidation inhibition assay. In another study, the methanolic extract of the seeds showed an average activity against DPPH with an inhibition value of 75% at 500 µL/mL [ 83 ]. According to these authors, this activity is probably due to the presence of polyphenols and cannabinoids, known for their strong antioxidant capacity [ 164 , 165 ]. Moreover, other seed metabolites, such as lignamides [ 156 ] and proteins [ 157 ], exhibited average antioxidant activities. However, they are lower than the results obtained with polyphenols and cannabinoids. Considering the high amount of polyphenols in the leaves, the extracts obtained from these plant parts have generally shown strong antioxidant activity [ 87 , 155 ]. For example, the hydroalcoholic extract of the leaves showed a DPPH IC 50 value of 2.7 mg/mL [ 155 ]. This value is relatively lower than the result obtained in the seed DPPH assay, which suggests a higher antioxidant potential. Moreover, the aerial essential parts showed also excellent antioxidant activity, with values near to the positive control [ 163 ]. As previously mentioned, polyphenols are generally considered as the main group of antioxidant molecules that work through different mechanisms, such as the suppression of free radicals that initiate oxidative damage and inhibit the oxidation process, via chelation of catalytic metals or metal ions and the inhibition of lipoxygenase [ 166 , 167 ]. Furthermore, some volatile terpenes exhibits potent antioxidant and anti-free-radical properties [ 168 ]. This can explain the antioxidant activity of the essential oils of the aerial parts proven through different tests.

4.4.2. Antimicrobial Activity

An antimicrobial is a molecule with microbicidal (kills microorganisms) or microbiostatic (slow the microbial growth and/or development) activity. These substances have different names depending on the type of targeted microorganism, such as antibacterial (for bacteria), antifungal (for fungi), antiviral (for virus) or antiparasitic (for parasites). The antimicrobial activity of essential oils and organic extracts of different parts of Cannabis sativa L. against several microorganisms has been reported. However, the degree of antimicrobial activity varies from cultivar to cultivar [ 169 ], as well as according to the part of the plant used, the extraction method and type of extract. The seed hydroalcoholic extract was evaluated against Gram-positive and -negative bacteria, namely Staphylococcus aureus , Escherichia coli , Salmonella typhimurium , Enterobacter aerogenes , Enterococcus faecalis , Lacticaseibacillus paracasei , Limosilactobacillus reuteri , Levilactobacillus brevis , Lactiplantibacillus plantarum , Bifidobacterium bifidum , Bifidobacterium longum and Bifidobacterium breve , and the obtained results showed low antibacterial activity with MIC values superior to 1 mg/mL [ 158 ]. Regarding antibacterial tests on Cannabis leaves, a study published by Anjum [ 159 ] compared the efficacy of four extracts obtained with acetone, chloroform, ethanol and water against three bacterial strains, namely Escherichia coli , Staphylococcus aureus and Pseudomonas aeruginosa . The results showed similarities between the four extracts with values near to 19 mm. Moreover, the results published by Manosroi, et al. [ 155 ] showed the effect of ethanolic extract as an antibacterial agent against Staphylococcus mutans with an inhibition diameter of 1.33 ± 0.58 mm. The essential oils of the aerial parts were evaluated as well for their antibacterial activity. The volatile terpenes of Cannabis exerted diverse activity intensities according to the targeted bacterial strains, where the weakest antibacterial activity was observed against Helicobacter pylori and Klebsiella pneumonia strains, with MICs values of 64 and 38 mg/mL, respectively. Moreover, a weak antibacterial activity was observed against Micrococcus luteus and Staphylococcus aureus , with an MIC of 4.7 mg/mL for both strains, whereas a more moderate inhibitory activity was observed against Escherichia coli , Pseudomonas aeruginosa and Bacillus subtilis , with an MIC of 1.2 mg/mL [ 163 ]. These results are explained by the fact that volatile terpenes are known to be strong antibacterial compounds, according to many biological investigations [ 170 , 171 , 172 ].

Concerning the antifungal activity, Anjum [ 159 ] compared the efficacy of four Cannabis leaf extracts, obtained with acetone, chloroform, ethanol and water, against two fungi, namely Aspergillus niger and Fusarium spp. The extracts showed similar results with inhibition diameter values ranging between 20.6 and 23 mm for Aspergillus niger , and 18.3 to 24.3 mm for Fusarium spp. In another studies, the acetone extract of Cannabis flowers showed a significant effect on the growth of Aspergillus favus , the flower hydroalcoholic extract led to the inhibition of 36% of fungi mycelium at concentration of 7.2 mg of dry matter per mL of culture medium [ 161 ]. Moreover, aerial part essential oils showed interesting antifungal potential. Nafis, et al. [ 163 ] reported an MIC value of 9.5 mg/mL against four fungi species, namely Candida albicans , Candida glabrata , Candida krusei and Candida parapsilosis . However, Zengin, et al. [ 94 ] discovered a weak antifungal activity against a group of clinically relevant and multidrug-resistant microorganisms belonging to Candida spp. and Malassezia spp., with essential oil MIC values superior to 12.460 µg/mL.

4.4.3. Insecticidal Activity

Insecticides are active substances with the property of killing insects, their larvae and/or eggs. Insecticides act either by contact or after penetration into the digestive tract or into the respiratory system. Essential oils from inflorescences were evaluated for their mosquitocidal activities on larvae and pupae of two main malaria vectors known as Anopheles gambiae and Anopheles stephensi . The results showed that Cannabis inflorescence essential oils showed toxicity against mosquitoes with LC 50 values of 73.5 to 78.8 ppm for Anopheles stephensi larvae, and 20.13 to 67.19 ppm for pupae of Anopheles gambiae . Their natural origin and volatile propriety makes the use of these essential oils very attractive for the formulation of stable and safe biocontrol products [ 162 ]. The cholinesterase inhibition activity had also been reported in the literature. Furthermore, a molecule purified from the ethanolic extract of Cannabis seeds, namely 3,3′-demethyl-heliotropamide, showed a moderate activity with an IC 50 value of 46.2 µm [ 156 ], whereas the aerial part essential oils showed stronger activity against butyryl-choline esterase, with a concentration of 3.4 mg GALAE/g oil [ 94 ].

4.4.4. Anticoagulant Activity

It has been suggested that plants with anticoagulant activities act as herbal remedies that could lead to the discovery of new therapeutic agents to treat thrombosis-related diseases. Blood clotting studies were conducted to determine the possible antiprothrombotic effect of Cannabis leaf metabolites, with three main cannabinoids, THC, CBD and CBN, targeted. The in vitro effect of Cannabis extract on thrombin activity was evaluated by Coetzee, et al. [ 160 ]. In their publication, two cannabinoids, namely THC and CBN, showed interesting IC 50 values. The highest activity was obtained by THC, with a value of 1.79 mg/mL, whereas CBN showed weaker activity suggested by high IC 50 value. However, this study also featured an in vivo test applied on obese rats in order to determine the clotting times. As a result, the Cannabis-treated rats showed an efficiency of 50% with clotting two times higher than the control groups, thus proving that cannabinoids may have a good anticoagulant activity.

4.4.5. Antidiabetic Activity

Diabetes is a chronic, progressive and complex metabolic disorder characterized by abnormally high blood glucose levels. This condition is also known as hyperglycemia. Worldwide, approximately 90% of affected patients are non-insulin-dependent, classified as type 2 diabetes [ 173 ]. According to the World Health Organization (WHO), diabetes is a chronic disease that occurs when the pancreas does not produce enough insulin, or the body does not properly use the produced insulin. Insulin is a hormone that regulates the concentration of sugar in the blood. Antidiabetic activity is generally evaluated by the quantification of α-amylase inhibition [ 174 ]. This enzyme is generally produced in the pancreas and plays a key enzyme role in the increase in blood sugar by breaking down dietary carbohydrates, such as starch, into simple monosaccharides in the digestive system, followed by the further α-glucosidase degradation into glucose which, upon absorption, enters in the bloodstream. Therefore, inhibition of the enzymes α-amylase and α-glucosidase can suppress carbohydrate digestion, delay glucose absorption and, consequently, reduce blood glucose levels [ 175 ]. The study published by Zengin, et al. [ 94 ] proved that Cannabis aerial part essential oils exhibited antidiabetic properties against the α-glucosidase enzyme with a value of 3.77 mmol ACAE/g oil. This essential oil has also been evaluated against α-amylase but showed no significant result.

4.4.6. Anticancer Activity

Cancer remains a major cause of morbidity and mortality worldwide. It is currently treated using classical approaches such as surgery, chemotherapy and radiotherapy. The toxic side effects associated with chemotherapy and radiotherapy often lead to adverse health effects. This explains the huge need for new drugs, safer to use with less side effects. Experimentally, several Cannabis-derived compounds demonstrated conclusive efficacy in vitro and in vivo on a wide range of cancer cell lines, including breast [ 176 ], prostate [ 177 ], cervix [ 178 ], brain [ 179 ], colon [ 180 ] and leukemia/lymphoma [ 181 ]. A number of in vitro and in vivo studies have demonstrated the effects of phytocannabinoids on tumor progression. These studies suggest that specific cannabinoids such as Δ9-THC and CBD induce apoptosis and inhibit proliferation in various cancer cell lines at concentrations ranging between 5 and 65 µm [ 176 , 182 , 183 , 184 , 185 , 186 , 187 ]. Moreover, combination of certain phytocannabinoids improved the anticancer activity of Cannabis preparations; for example, Armstrong, et al. [ 182 ] revealed that the combination of CBD and Δ9 -THC exhibited a stronger melanoma cell mortality in comparison with Δ9-THC alone. In general, phytochemicals in the Cannabis plant, and especially cannabinoids, are non-selective in their functions and limited in their differential activity on cancer cells with normal cells. Therefore, researchers show interest in isolating bioactive phytochemicals from Cannabis with potent anticancer properties and generating lead compounds based on the natural backbone of a molecule as a synthetic approach.

4.4.7. Anti-Inflammatory and Analgesic Activities

Many compounds of Cannabis proved to have strong anti-inflammatory activity. The Cannabis seeds showed an inflammation-reducing capacity, especially on primary human monocytes treated with LPS. The results proved that the compounds of these seeds decreased the respective expression and secretion of IL-6 genes and TNF-α. Additionally, cannabinoids proved to be strong anti-inflammatory agents; in fact, they can suppress the production of pro-inflammatory cytokines and chemokines and may have therapeutic applications in health conditions underlying inflammatory components [ 188 , 189 ]. Analgesic action is defined as any procedure whose principle of activity is to reduce pain. This can be not only a drug but also any other method aimed at achieving analgesia, i.e., the abolition of the sensation of pain [ 190 ]. Clinical and experimental studies showed that Cannabis-derived compounds act as analgesic agents. However, the effectiveness of each product is variable and depends on the administration mode. With opioids being the only therapy for severe pain, the analgesic capacity of cannabinoids could provide a much-needed alternative to opioids [ 191 ]. Furthermore, cannabinoids act synergistically with opioids and act as opioid-sparing agents, allowing lower doses and fewer side effects of chronic opioid treatment [ 192 ]. Thus, the rational use of Cannabis-based medicines deserves to be seriously considered to alleviate patients’ suffering from severe pain.

4.4.8. Neuroprotective Activity

The terpenes and cannabinoids of Cannabis proved also to have neuroprotective proprieties. The neuroprotective effects of 17 compounds present in the aerial parts of Cannabis sativa L. were evaluated in PC-12 cells including p-hydroxybenzaldehyde, (e)-methyl p-hydroxycinnamate and ferulic acid—which showed additional protective effects against H 2 O 2 -induced damage [ 193 ]. Furthermore, di Giacomo, et al. [ 147 ] reported the neuroprotective and neuromodulatory effects induced by cannabidiol and cannabigerol in rat Hypo-E22 cells and isolated hypothalamus, whereas Landucci, et al. [ 194 ] proved that appropriate concentrations of CBD or CBD/THC ratios can represent a valid therapeutic intervention in the treatment of post-ischemic neuronal death. In another study, Esposito, et al. [ 195 ] highlighted the importance of CBD as a promising new drug able to reduce neuroinflammatory responses evoked by β-amyloid. Furthermore, the study published by Perez, et al. [ 196 ] described the neuronal counting of both motor and sensory neurons after CBD treatment using immunohistochemical analysis. The obtained results showed an increase by 30% of synaptic preservation on the spinal cord for the CBD-treated group, suggesting an average neuroprotective effect.

4.4.9. Antiepileptic and Anticonvulsant Activities

Despite being well-known for its psychoactive proprieties, Cannabis sativa L. has been investigated for additional effects on the central nervous system. A recently published study showed that CBD has a high efficacy in epilepsy with hippocampal focus than with the extrahippocampal amygdala and parvalbumin, implying a protective role in regulating hippocampal seizures and neurotoxicity at a juvenile age [ 197 ]. The efficiency of CBD has been further replicated in human populations, including adolescents and young adults with severe childhood-onset epilepsy [ 198 ]. In a 12-week open-label trial, a group of patients aged between 1 and 30 years were treated with 25 and 50 mg/kg of CBD. The treated groups showed a reduction in epileptic attack severity by 36.5%. Preclinical research has also attempted a more chronic elucidation of the efficacy of CBD as an anticonvulsant. Using the PTZ model of epilepsy, it was found that a reduction in seizure activity could be achieved with varying doses between 20 and 50 mg/kg of CBD over a 28-day treatment period [ 199 ]. Other cannabinoids, such as cannabigerol, cannabidivarin, cannabichromene, δ 9 -tetrahydrocannabinolic acid and tetrahydrocannabivarin, showed efficacy in models of Huntington’s disease and epilepsy. It is important to note that these phytocannabinoids and their combinations are warranted in a range of other neurodegenerative disorders such as Parkinson’s [ 200 ].

4.4.10. Dermocosmetic Activity

Lipids in human skin play a very important role in preserving the structure of the dermis, protecting it from dehydration. However, during menopause, hormonal changes negatively affect the skin’s balance, making it more prone to developing dryness [ 201 ]. The underlying tissues, such as subcutaneous adipose tissue and muscles, undergo atrophy due to the overproduction of some enzymes—among them tyrosinase, elastase and collagenase. Tyrosinase is mainly responsible for the production of melanin on the skin, whereas collagenase and elastase target the skin structural proteins collagen and elastin and degrade them, respectively. The results published by Manosroi, et al. [ 155 ] showed that leaf extracts have anti-tyrosinase activity with an IC 50 value of 0.07 ± 0.06 mg/mL, which suggest a strong tyrosinase inhibitory activity. Similarly, Zagórska-Dziok et al. [ 87 ] showed respective collagenase and elastase inhibitory activities of 80 and 30% at 1000 µg/mL. These activities are mainly due to the presence of polyphenols and cannabinoids.

4.5. Drugs Based on Cannabis sativa L.

Due to the importance of the biological evaluation’s findings, several Cannabis-based commercial pharmaceuticals have been produced. These products have many biological proprieties and were produced to treat a wide range of conditions. The first reported product has been commercialized under the name “marinol”. This drug was developed 40 years ago, in 1985, by an American pharmaceutical company, and used dronabinol and synthetic THC as active agents [ 202 ]. Likewise, “syndros” is another drug marketed in the USA in 2016 and it contains the same active ingredients [ 202 ]. Both drugs are indicated for the treatment of severe nausea and vomiting related to cancer chemotherapy and AIDS-anorexia associated with weight loss. Another active compound known as “nabilone” is used as ingredient of two drugs, “casamet” and “canemes” [ 203 ]. The “nabilone” is a synthetic analogue of THC approved by the U.S. Food and Drug Administration (U.S. FDA) for the treatment of chemotherapy and AIDS symptoms [ 204 ]. Two other cannabinoids, namely “CBD” and “THC”, are also used in two drugs commercialized under the name of Bourneville and used against two types of severe epilepsy (Lennox–Gastaut syndrome and Dravet syndrome) [ 205 ], whereas Sativex, generally known as “nabiximols”, is used to alleviate muscle spasms in multiple sclerosis disease.

5. Conclusions

This manuscript has reviewed and analyzed the historical, botanical, ethnopharmacological, chemical, bioinformatics and biological knowledge of Cannabis sativa from the earliest human communities to current medical applications, with a critical analysis of the multiple potential applications of cannabinoids in the contemporary scientific context.

At present, more than 545 phytochemicals have been described in the different parts of the Cannabis plant. The most represented metabolite class is the phytocannabinoids and they exhibit enormous structural diversity and bioactivities. Cannabis sativa is found in a wide variety of forms and environments on all continents and its pharmacological properties seem to go far beyond psychotic effects, with the ability to address needs such as the treatment and relief of many symptoms and diseases.

Furthermore, the relaxation of regulatory standards for therapeutic Cannabis and the conduct of more controlled clinical trials suggests that the Cannabis sativa plant has interesting therapeutic potential as an antiemetic, appetite stimulant in debilitating diseases (cancer and AIDS), analgesic, as well as in the treatment of multiple sclerosis, spinal cord injury, Tourette syndrome, epilepsy and glaucoma. Further clinical research is needed to investigate the potential therapeutic uses of this plant in specific medical conditions. Scientifically designed trials will help establish which of the cannabinoids produce the various beneficial effects described, or whether these are the result of a combination of cannabinoids. The research would also help to better characterize the adverse effects of each cannabinoid.

Acknowledgments

We would like to thank COST Action 18101 SOURDOMICS—Sourdough biotechnology network towards novel, healthier and sustainable food and bioprocesses ( https://sourdomics.com/ ; https://www.cost.eu/actions/CA18101/ , accessed on 7 February 2023), where the author N.E.A. is member of the working groups 3, 4 and 7, and the author J.M.R. is the Chair and Grant Holder Scientific Representative and is supported by COST (European Cooperation in Science and Technology) ( https://www.cost.eu/ , accessed on 7 February 2023). COST is a funding agency for research and innovation networks. Regarding the author J.M.R., he is financially supported by LA/P/0045/2020 (ALiCE) and UIDB/00511/2020-UIDP/00511/2020 (LEPABE) funded by national funds through FCT/MCTES (PIDDAC).

Funding Statement

“Agence National des Plantes Médicinales et Aromatiques-Taounate, ANPMA” Morocco and University Abdelmalek Essaadi-Tetouan Morocco (VPMA4).

Author Contributions

Conceptualization: N.E.A. and S.H.; Data collection: S.H., A.Y.B. and H.M.; Writing—original draft preparation: S.H. and H.M.; Writing—review and editing: H.M., J.M.R. and N.E.A.; Supervision: N.E.A. and J.M.R.; Funding acquisition: N.E.A. and J.M.R. All authors have read and agreed to the published version of the manuscript.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Data availability statement, conflicts of interest.

The authors declare no conflict of interest.

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Computer Science > Computation and Language

Title: realm: reference resolution as language modeling.

Abstract: Reference resolution is an important problem, one that is essential to understand and successfully handle context of different kinds. This context includes both previous turns and context that pertains to non-conversational entities, such as entities on the user's screen or those running in the background. While LLMs have been shown to be extremely powerful for a variety of tasks, their use in reference resolution, particularly for non-conversational entities, remains underutilized. This paper demonstrates how LLMs can be used to create an extremely effective system to resolve references of various types, by showing how reference resolution can be converted into a language modeling problem, despite involving forms of entities like those on screen that are not traditionally conducive to being reduced to a text-only modality. We demonstrate large improvements over an existing system with similar functionality across different types of references, with our smallest model obtaining absolute gains of over 5% for on-screen references. We also benchmark against GPT-3.5 and GPT-4, with our smallest model achieving performance comparable to that of GPT-4, and our larger models substantially outperforming it.

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