Avocado Genome Elucidated

McDonnell Boehnen Hulbert & Berghoff LLP
Contact

McDonnell Boehnen Hulbert & Berghoff LLP

Persea_americana_fruit_2 The avocado, having gained popularity (at least in the U.S.) as a convenient (and delicious) vehicle for consuming otherwise not particularly healthful corn chips, has more recently been hailed as a "superfood" when consumed in diets as a healthy form of fat. Recently, the avocado joined the ranks of other plants important for human nutrition (see, for example, "Genomic Sequence of Strawberry Determined"; "Genetic Analyses of Sweet Potato Genome Sheds Light on Speciation and Global Dispersion Patterns"; "Durum Wheat Genome Revealed"; and "Genetic Assessment of Squash Genomes in Related Species") in having the sequence of its genome determined, with concomitant explication of its evolutionary history and relationship to other members of the plant kingdom.

This achievement was reported in the Proceedings of the National Academy of Sciences USA, in a paper entitled "The avocado genome informs deep angiosperm phylogeny, highlights introgressive hybridization, and reveals pathogen-influenced gene space adaptation" (https://doi.org/10.1073/pnas.1822129116) by an international group of researchers.* The avocado, Persea americana, is a member of the Lauraceae family, otherwise known for the spices cinnamon, bay leaves, and sassafras (gumbo filé). Avocadoes anciently diverged from other angiosperms, and their magnoliid clade comprises 11,000 total species, which the authors assert is "minuscule in comparison to the dominant eudicot and monocot flowering plant lineages, comprising about 285,000 species combined." Conventional genetic analyses had not established the avocados' phylogenetic position relative to eudicots and monocots.

The predominant avocado is the Mexican avocado, P. americana var. drymifolia, comprising three varieties: the Mexican, Guatemalan, and West Indian varieties. Familiar to many afficianados is the Hass cultivar, which is a hybrid between the Guatemalan and Mexican races and is typically grafted onto Mexican race rootstock. Worldwide, 90% of cultivated avocado corresponds to the cultivar Hass; surprisingly, the Hass cultivar is of recent (20th century) origin.

The authors sequenced Hass and Mexican cultivars as reference genomes, and resequenced three other cutivars (vars. drymifolia, guatemalensis, and americana), as well as wild avocados of the West Indian variety (P. america1na var. costaricensis), and Persea shiedeana (the edible coyo), a species that is relatively closely related to P. americana. The avocado genome comprises 12 chromosomes, and these authors report that the genome size of P. americana var. drymifolia is 920 Mb in size, while the P. americana Hass cultivar has a 912.6 Mb estimated genome size. Analysis of protein coding sequences was based on a comparison with protein sequences from the following angiosperm species: Sorghum bicolor (33,032 proteins), Vitis vinifera (26,343 proteins), Solanum lycopersicum (34,727 proteins), and Arabidopsis thaliana (27,416 proteins). These authors reported an estimated number of protein-coding genes in each genome: 22,441 protein-coding genes from the Mexican genome and 24,616 protein-coding genes from the Hass cultivar genome.

Turning to genome structure, the authors used principal component analysis of genome-wide single nucleotide polymorphisms (SNPs) to show relative uniformity in Costa Rican/West Indian/Guatemalan cultivars but strong heterogeneity within the Mexican subpopulation. The paper reported 5,050 SNP markers, distributed as follows:

Table

Figure 2AResults of these analyses of the Hass cultivar were intermediate between the Guatemalan and Mexican subpopulations, which was in agreement with the hybrid nature of this variety. Also consistent with the recent appearance of the Hass cultivar, its genome showed significant (39%) introgression of DNA having Guatamalan variety structure into a Mexican genomic background. Haas cultivar Chromosome 4 illustrates these analyses, demonstrating that a "huge" Guatemalan block, which could encompass an entire chromosomal arm, is present in the Hass genome. Figure 2A (at right) in the article illustrates a 22 megabase (Mb) stretch of DNA on chromosome 4 exemplifyng genomic introgression from the Guatemalan avocado race (var. guatemalensis) into a Mexican (var. drymifolia) genetic background.

The authors further report that the relatively long length of this Guatemalan-derived block is uninterrupted by recombination, reflecting the extremely recent hybrid origin of the cultivar.

Further, genomic analysis revealed evidence of two "ancient" polyploidy events, which the authors report are lineage-specific and both postdate the divergence of the avocado from common ancestry with other angiosperm species. Avocado divergence from other species appears to have occurred from 7.4 to 3.8 million years ago. Unfortunately, these relationships could not be unambiguously resolved because the outcome varied with the way the analysis was performed. Based on comparison of protein sequences, for example, avocado was resolved as a sister to monocots plus eudicots (i.e., branching occurred before their divergence from each other), whereas from analysis of coding sequences avocado was placed as sister to monocots only.

The authors also reported the phenotypic effects of genome structure, specifically the occurrence of tandem duplications, on gene expression related to potentially important metabolic responses. Enrichment of tandem duplications was found in chromosomal regions that the authors contend could be related to recent adaptation against fungal pathogens. On the other hand, ancient polyploid duplicates were enriched with "transcriptional regulatory functions reflective of core physiological and developmental processes" (specifically, "among 2,433 total polyploid duplicates, regulation of transcription, DNA-templated was significantly overrepresented by 352 genes"). Transcriptional activity in tandem duplicates was found to increase following anthracnose infection, the authors positing that some of the up-regulated genes could be related to defense responses. These included genes involved in phenylpropanoid biosynthesis and closely related pathways are significantly enriched among tandem duplicates. The authors further state:

This functional enrichment in a long-lived tree may have evolved in response to pathogen infection, including Colletotrichum (anthracnose) and Phytophthora cinnamomi (avocado root rot), both of which are reported to activate the phenylpropanoid biosynthetic pathways in avocado. . . . Several . . . functional enrichments among avocado tandems (for example, 1,3-beta-D-glucan synthase activity and regulation of cell shape) relate to callose synthase activity, a recently discovered avocado defense mechanism against P. cinnamomi. Other significantly enriched [metabolic pathways] include phenylpropanoid metabolic process, lignin biosynthetic process, and UDP-glycosyltransferase activity, categories directly or closely related to phenylpropanoid biosynthesis. The lignin functional enrichment, for example, includes diverse tandemly duplicated genes involved in many pathway-interrelated processes, including homologs of both biosynthetic and regulatory genes encoding hydroxycinnamoyl-CoA shikimate/quinate hydroxycinnamoyl transferase (HCT), cinnamyl alcohol dehydrogenase 5 (CAD5), laccase 17 (LAC17), caffeate o-methyltransferase 1 (COMT1), peroxidase 52 (PRX52), NAC domain containing protein 12 (NAC012), and NAC secondary wall thickening promoting factor 1 (NST1). As could be expected from the above, the [metabolic pathways] involved in defense response and defense response to fungus are significantly enriched among tandem duplicates, as has been discovered for other plant genomes, and involving many different gene families and responses. Tandem O-methyltransferases homologous to COMT1 may also contribute to synthesis of the phenylpropanoid derivative and insecticide estragole, which is largely responsible for the anise-like leaf scent and fruit taste of many avocado cultivars, particularly of the Mexican race. Another relevant enriched . . . category among tandems is ethylene-activated signaling pathway, which annotates many different transcription factor duplicates. Ethylene signaling factors such as ERF1 (represented by 2 homologs) are heavily involved in pathogen-induced responses, including to infection by Colletotrichum and other necrotrophic fungi. Also identified are 3 homologs of EIN3, a transcription factor that initiates downstream ethylene responses, including fruit ripening. Avocado fruit matures on the tree in a process that involves ethylene synthesis and signaling, while it does not ripen until harvested—a desirable trait that allows growers to delay harvesting for several months.

The authors characterize tandem duplicates as "evolutionarily recent 'tuning knobs' in the genome adaptive landscape[]" of the avocado species:

[M]ost tandem duplicates in the genome are expected to be of more recent origin, having been generated by ongoing gene birth–death–innovation processes that operate in all eukaryotic genomes. As such, sub- or neofunctionalized tandem duplicates that survive the usual fate of duplicated genes—pseudogenization—should be enriched in functions that fine-tune a given species' recent selective environment. In the case of avocado, response to fungal pathogens is precisely reflected in its tandemly duplicated gene complement.

The authors conclude their report with a concise explanation of its significance:

Our genomes of Mexican and Hass avocados provide the requisite resources for genome-wide association studies to identify important traits among natural avocado genetic diversity present in Mesoamerica, to develop genome-assisted breeding and genetic modification efforts crucial for the improvement of this long-life-cycle crop, to fight threatening avocado diseases, and to optimize growth and desirable phenotypic traits.

* Martha Rendón-Anayaa,b,1,
Enrique Ibarra-Laclettea,c,1,
Alfonso Méndez-Bravoa,d,
Tianying Lane,
Chunfang Zhengf,
Lorenzo Carretero-Pauletg,
Claudia Anahí Perez-Torresa,c,
Alejandra Chacón-Lópeza,
Gustavo Hernandez-Guzmána,h,i,
Tien-Hao Change,
Kimberly M. Farre,
Brad Barbazukj,
Srikar Chamalak,
Marek Mutwill,
Devendra Shivharel,
David Alvarez-Poncem,
Neena Mittern,
Alice Haywardn,
Stephen Fletchern,
Julio Rozaso,p,
Alejandro Sánchez Graciao,p,
David Kuhnq,
Alejandro F. Barrientos-Priegor,
Jarkko Salojärvil,
Pablo Librados,t,
David Sankofff,
Alfredo Herrera-Estrellaa,
Victor A. Alberte,l,2, and
Luis Herrera-Estrellaa,u,2

aUnidad de Genomica Avanzada/Langebio, Centro de Investigación y de Estudios Avanzados, Irapuato 36821, México; bDepartment of Plant Biology, Uppsala BioCenter, Swedish University of Agricultural Sciences, SE-750 07 Uppsala, Sweden; cRed de Estudios Moleculares Avanzados, Instituto de Ecología A.C., 91070 Xalapa, México; dEscuela Nacional de Estudios Superiores, Laboratorio Nacional de Análisis y Síntesis Ecológica, Universidad Nacional Autónoma de México, 58190 Morelia, México; eDepartment of Biological Sciences, University at Buffalo, Buffalo, NY 14260; fDepartment of Mathematics and Statistics, University of Ottawa, Ottawa, ON, Canada K1N 6N5; gCenter for Plant Systems Biology, Vlaams Instituut voor Biotechnologie (VIB), University of Ghent, 9052 Ghent, Belgium; hDepartamento de Alimentos, Universidad de Guanajuato, 36500 Irapuato, México; iDivisión de Ciencias de la Vida, Universidad de Guanajuato, 36500 Irapuato, México; jDepartment of Biology, University of Florida, Gainesville, FL 32611; kDepartment of Pathology, Immunology and Laboratory Medicine, University of Florida, Gainesville, FL 32610; lSchool of Biological Sciences, Nanyang Technological University, Singapore 637551; mDepartment of Biology, University of Nevada, Reno, NV 89557; nCentre for Horticultural Science, Queensland Alliance for Agriculture and Food Innovation, The University of Queensland, St. Lucia, QLD 4072, Australia; oDepartament de Genètica, Microbiologia i Estadística, Universitat de Barcelona, 08007 Barcelona, Spain; pInstitut de Recerca de la Biodiversitat, Universitat de Barcelona, 08007 Barcelona, Spain; qSubtropical Horticulture Research Station, Agricultural Research Service, US Department of Agriculture, Miami, FL 33158; rPosgrado en Horticultura, Departamento de Fitotecnia, Universidad Autónoma Chapingo, 56230 Texcoco, México; sCentre for GeoGenetics, Natural History Museum of Denmark, 1017 Copenhagen, Denmark; tLaboratoire d'Anthropobiologie Moléculaire et d'Imagerie de Synthèse, CNRS Unité Mixte de Recherche 5288, Université de Toulouse, Université Paul Sabatier, 31330 Toulouse, France; uDepartment of Plant and Soil Science, Texas Tech University, Lubbock, TX 79409

Image of Avocados (Persea americana) by B.navez, from the Wikimedia Commons under the Creative Commons Attribution-Share Alike 3.0 Unported, 2.5 Generic, 2.0 Generic and 1.0 Generic license.

DISCLAIMER: Because of the generality of this update, the information provided herein may not be applicable in all situations and should not be acted upon without specific legal advice based on particular situations.

© McDonnell Boehnen Hulbert & Berghoff LLP | Attorney Advertising

Written by:

McDonnell Boehnen Hulbert & Berghoff LLP
Contact
more
less

McDonnell Boehnen Hulbert & Berghoff LLP on:

Readers' Choice 2017
Reporters on Deadline

"My best business intelligence, in one easy email…"

Your first step to building a free, personalized, morning email brief covering pertinent authors and topics on JD Supra:
*By using the service, you signify your acceptance of JD Supra's Privacy Policy.
Custom Email Digest
- hide

JD Supra Privacy Policy

Updated: May 25, 2018:

JD Supra is a legal publishing service that connects experts and their content with broader audiences of professionals, journalists and associations.

This Privacy Policy describes how JD Supra, LLC ("JD Supra" or "we," "us," or "our") collects, uses and shares personal data collected from visitors to our website (located at www.jdsupra.com) (our "Website") who view only publicly-available content as well as subscribers to our services (such as our email digests or author tools)(our "Services"). By using our Website and registering for one of our Services, you are agreeing to the terms of this Privacy Policy.

Please note that if you subscribe to one of our Services, you can make choices about how we collect, use and share your information through our Privacy Center under the "My Account" dashboard (available if you are logged into your JD Supra account).

Collection of Information

Registration Information. When you register with JD Supra for our Website and Services, either as an author or as a subscriber, you will be asked to provide identifying information to create your JD Supra account ("Registration Data"), such as your:

  • Email
  • First Name
  • Last Name
  • Company Name
  • Company Industry
  • Title
  • Country

Other Information: We also collect other information you may voluntarily provide. This may include content you provide for publication. We may also receive your communications with others through our Website and Services (such as contacting an author through our Website) or communications directly with us (such as through email, feedback or other forms or social media). If you are a subscribed user, we will also collect your user preferences, such as the types of articles you would like to read.

Information from third parties (such as, from your employer or LinkedIn): We may also receive information about you from third party sources. For example, your employer may provide your information to us, such as in connection with an article submitted by your employer for publication. If you choose to use LinkedIn to subscribe to our Website and Services, we also collect information related to your LinkedIn account and profile.

Your interactions with our Website and Services: As is true of most websites, we gather certain information automatically. This information includes IP addresses, browser type, Internet service provider (ISP), referring/exit pages, operating system, date/time stamp and clickstream data. We use this information to analyze trends, to administer the Website and our Services, to improve the content and performance of our Website and Services, and to track users' movements around the site. We may also link this automatically-collected data to personal information, for example, to inform authors about who has read their articles. Some of this data is collected through information sent by your web browser. We also use cookies and other tracking technologies to collect this information. To learn more about cookies and other tracking technologies that JD Supra may use on our Website and Services please see our "Cookies Guide" page.

How do we use this information?

We use the information and data we collect principally in order to provide our Website and Services. More specifically, we may use your personal information to:

  • Operate our Website and Services and publish content;
  • Distribute content to you in accordance with your preferences as well as to provide other notifications to you (for example, updates about our policies and terms);
  • Measure readership and usage of the Website and Services;
  • Communicate with you regarding your questions and requests;
  • Authenticate users and to provide for the safety and security of our Website and Services;
  • Conduct research and similar activities to improve our Website and Services; and
  • Comply with our legal and regulatory responsibilities and to enforce our rights.

How is your information shared?

  • Content and other public information (such as an author profile) is shared on our Website and Services, including via email digests and social media feeds, and is accessible to the general public.
  • If you choose to use our Website and Services to communicate directly with a company or individual, such communication may be shared accordingly.
  • Readership information is provided to publishing law firms and authors of content to give them insight into their readership and to help them to improve their content.
  • Our Website may offer you the opportunity to share information through our Website, such as through Facebook's "Like" or Twitter's "Tweet" button. We offer this functionality to help generate interest in our Website and content and to permit you to recommend content to your contacts. You should be aware that sharing through such functionality may result in information being collected by the applicable social media network and possibly being made publicly available (for example, through a search engine). Any such information collection would be subject to such third party social media network's privacy policy.
  • Your information may also be shared to parties who support our business, such as professional advisors as well as web-hosting providers, analytics providers and other information technology providers.
  • Any court, governmental authority, law enforcement agency or other third party where we believe disclosure is necessary to comply with a legal or regulatory obligation, or otherwise to protect our rights, the rights of any third party or individuals' personal safety, or to detect, prevent, or otherwise address fraud, security or safety issues.
  • To our affiliated entities and in connection with the sale, assignment or other transfer of our company or our business.

How We Protect Your Information

JD Supra takes reasonable and appropriate precautions to insure that user information is protected from loss, misuse and unauthorized access, disclosure, alteration and destruction. We restrict access to user information to those individuals who reasonably need access to perform their job functions, such as our third party email service, customer service personnel and technical staff. You should keep in mind that no Internet transmission is ever 100% secure or error-free. Where you use log-in credentials (usernames, passwords) on our Website, please remember that it is your responsibility to safeguard them. If you believe that your log-in credentials have been compromised, please contact us at privacy@jdsupra.com.

Children's Information

Our Website and Services are not directed at children under the age of 16 and we do not knowingly collect personal information from children under the age of 16 through our Website and/or Services. If you have reason to believe that a child under the age of 16 has provided personal information to us, please contact us, and we will endeavor to delete that information from our databases.

Links to Other Websites

Our Website and Services may contain links to other websites. The operators of such other websites may collect information about you, including through cookies or other technologies. If you are using our Website or Services and click a link to another site, you will leave our Website and this Policy will not apply to your use of and activity on those other sites. We encourage you to read the legal notices posted on those sites, including their privacy policies. We are not responsible for the data collection and use practices of such other sites. This Policy applies solely to the information collected in connection with your use of our Website and Services and does not apply to any practices conducted offline or in connection with any other websites.

Information for EU and Swiss Residents

JD Supra's principal place of business is in the United States. By subscribing to our website, you expressly consent to your information being processed in the United States.

  • Our Legal Basis for Processing: Generally, we rely on our legitimate interests in order to process your personal information. For example, we rely on this legal ground if we use your personal information to manage your Registration Data and administer our relationship with you; to deliver our Website and Services; understand and improve our Website and Services; report reader analytics to our authors; to personalize your experience on our Website and Services; and where necessary to protect or defend our or another's rights or property, or to detect, prevent, or otherwise address fraud, security, safety or privacy issues. Please see Article 6(1)(f) of the E.U. General Data Protection Regulation ("GDPR") In addition, there may be other situations where other grounds for processing may exist, such as where processing is a result of legal requirements (GDPR Article 6(1)(c)) or for reasons of public interest (GDPR Article 6(1)(e)). Please see the "Your Rights" section of this Privacy Policy immediately below for more information about how you may request that we limit or refrain from processing your personal information.
  • Your Rights
    • Right of Access/Portability: You can ask to review details about the information we hold about you and how that information has been used and disclosed. Note that we may request to verify your identification before fulfilling your request. You can also request that your personal information is provided to you in a commonly used electronic format so that you can share it with other organizations.
    • Right to Correct Information: You may ask that we make corrections to any information we hold, if you believe such correction to be necessary.
    • Right to Restrict Our Processing or Erasure of Information: You also have the right in certain circumstances to ask us to restrict processing of your personal information or to erase your personal information. Where you have consented to our use of your personal information, you can withdraw your consent at any time.

You can make a request to exercise any of these rights by emailing us at privacy@jdsupra.com or by writing to us at:

Privacy Officer
JD Supra, LLC
10 Liberty Ship Way, Suite 300
Sausalito, California 94965

You can also manage your profile and subscriptions through our Privacy Center under the "My Account" dashboard.

We will make all practical efforts to respect your wishes. There may be times, however, where we are not able to fulfill your request, for example, if applicable law prohibits our compliance. Please note that JD Supra does not use "automatic decision making" or "profiling" as those terms are defined in the GDPR.

  • Timeframe for retaining your personal information: We will retain your personal information in a form that identifies you only for as long as it serves the purpose(s) for which it was initially collected as stated in this Privacy Policy, or subsequently authorized. We may continue processing your personal information for longer periods, but only for the time and to the extent such processing reasonably serves the purposes of archiving in the public interest, journalism, literature and art, scientific or historical research and statistical analysis, and subject to the protection of this Privacy Policy. For example, if you are an author, your personal information may continue to be published in connection with your article indefinitely. When we have no ongoing legitimate business need to process your personal information, we will either delete or anonymize it, or, if this is not possible (for example, because your personal information has been stored in backup archives), then we will securely store your personal information and isolate it from any further processing until deletion is possible.
  • Onward Transfer to Third Parties: As noted in the "How We Share Your Data" Section above, JD Supra may share your information with third parties. When JD Supra discloses your personal information to third parties, we have ensured that such third parties have either certified under the EU-U.S. or Swiss Privacy Shield Framework and will process all personal data received from EU member states/Switzerland in reliance on the applicable Privacy Shield Framework or that they have been subjected to strict contractual provisions in their contract with us to guarantee an adequate level of data protection for your data.

California Privacy Rights

Pursuant to Section 1798.83 of the California Civil Code, our customers who are California residents have the right to request certain information regarding our disclosure of personal information to third parties for their direct marketing purposes.

You can make a request for this information by emailing us at privacy@jdsupra.com or by writing to us at:

Privacy Officer
JD Supra, LLC
10 Liberty Ship Way, Suite 300
Sausalito, California 94965

Some browsers have incorporated a Do Not Track (DNT) feature. These features, when turned on, send a signal that you prefer that the website you are visiting not collect and use data regarding your online searching and browsing activities. As there is not yet a common understanding on how to interpret the DNT signal, we currently do not respond to DNT signals on our site.

Access/Correct/Update/Delete Personal Information

For non-EU/Swiss residents, if you would like to know what personal information we have about you, you can send an e-mail to privacy@jdsupra.com. We will be in contact with you (by mail or otherwise) to verify your identity and provide you the information you request. We will respond within 30 days to your request for access to your personal information. In some cases, we may not be able to remove your personal information, in which case we will let you know if we are unable to do so and why. If you would like to correct or update your personal information, you can manage your profile and subscriptions through our Privacy Center under the "My Account" dashboard. If you would like to delete your account or remove your information from our Website and Services, send an e-mail to privacy@jdsupra.com.

Changes in Our Privacy Policy

We reserve the right to change this Privacy Policy at any time. Please refer to the date at the top of this page to determine when this Policy was last revised. Any changes to our Privacy Policy will become effective upon posting of the revised policy on the Website. By continuing to use our Website and Services following such changes, you will be deemed to have agreed to such changes.

Contacting JD Supra

If you have any questions about this Privacy Policy, the practices of this site, your dealings with our Website or Services, or if you would like to change any of the information you have provided to us, please contact us at: privacy@jdsupra.com.

JD Supra Cookie Guide

As with many websites, JD Supra's website (located at www.jdsupra.com) (our "Website") and our services (such as our email article digests)(our "Services") use a standard technology called a "cookie" and other similar technologies (such as, pixels and web beacons), which are small data files that are transferred to your computer when you use our Website and Services. These technologies automatically identify your browser whenever you interact with our Website and Services.

How We Use Cookies and Other Tracking Technologies

We use cookies and other tracking technologies to:

  1. Improve the user experience on our Website and Services;
  2. Store the authorization token that users receive when they login to the private areas of our Website. This token is specific to a user's login session and requires a valid username and password to obtain. It is required to access the user's profile information, subscriptions, and analytics;
  3. Track anonymous site usage; and
  4. Permit connectivity with social media networks to permit content sharing.

There are different types of cookies and other technologies used our Website, notably:

  • "Session cookies" - These cookies only last as long as your online session, and disappear from your computer or device when you close your browser (like Internet Explorer, Google Chrome or Safari).
  • "Persistent cookies" - These cookies stay on your computer or device after your browser has been closed and last for a time specified in the cookie. We use persistent cookies when we need to know who you are for more than one browsing session. For example, we use them to remember your preferences for the next time you visit.
  • "Web Beacons/Pixels" - Some of our web pages and emails may also contain small electronic images known as web beacons, clear GIFs or single-pixel GIFs. These images are placed on a web page or email and typically work in conjunction with cookies to collect data. We use these images to identify our users and user behavior, such as counting the number of users who have visited a web page or acted upon one of our email digests.

JD Supra Cookies. We place our own cookies on your computer to track certain information about you while you are using our Website and Services. For example, we place a session cookie on your computer each time you visit our Website. We use these cookies to allow you to log-in to your subscriber account. In addition, through these cookies we are able to collect information about how you use the Website, including what browser you may be using, your IP address, and the URL address you came from upon visiting our Website and the URL you next visit (even if those URLs are not on our Website). We also utilize email web beacons to monitor whether our emails are being delivered and read. We also use these tools to help deliver reader analytics to our authors to give them insight into their readership and help them to improve their content, so that it is most useful for our users.

Analytics/Performance Cookies. JD Supra also uses the following analytic tools to help us analyze the performance of our Website and Services as well as how visitors use our Website and Services:

  • HubSpot - For more information about HubSpot cookies, please visit legal.hubspot.com/privacy-policy.
  • New Relic - For more information on New Relic cookies, please visit www.newrelic.com/privacy.
  • Google Analytics - For more information on Google Analytics cookies, visit www.google.com/policies. To opt-out of being tracked by Google Analytics across all websites visit http://tools.google.com/dlpage/gaoptout. This will allow you to download and install a Google Analytics cookie-free web browser.

Facebook, Twitter and other Social Network Cookies. Our content pages allow you to share content appearing on our Website and Services to your social media accounts through the "Like," "Tweet," or similar buttons displayed on such pages. To accomplish this Service, we embed code that such third party social networks provide and that we do not control. These buttons know that you are logged in to your social network account and therefore such social networks could also know that you are viewing the JD Supra Website.

Controlling and Deleting Cookies

If you would like to change how a browser uses cookies, including blocking or deleting cookies from the JD Supra Website and Services you can do so by changing the settings in your web browser. To control cookies, most browsers allow you to either accept or reject all cookies, only accept certain types of cookies, or prompt you every time a site wishes to save a cookie. It's also easy to delete cookies that are already saved on your device by a browser.

The processes for controlling and deleting cookies vary depending on which browser you use. To find out how to do so with a particular browser, you can use your browser's "Help" function or alternatively, you can visit http://www.aboutcookies.org which explains, step-by-step, how to control and delete cookies in most browsers.

Updates to This Policy

We may update this cookie policy and our Privacy Policy from time-to-time, particularly as technology changes. You can always check this page for the latest version. We may also notify you of changes to our privacy policy by email.

Contacting JD Supra

If you have any questions about how we use cookies and other tracking technologies, please contact us at: privacy@jdsupra.com.

- hide

This website uses cookies to improve user experience, track anonymous site usage, store authorization tokens and permit sharing on social media networks. By continuing to browse this website you accept the use of cookies. Click here to read more about how we use cookies.