adulthood learning essay

Rethinking adult development

June 9, 2020

What is the special issue about?

We focus in this special issue of American Psychologist on how adulthood is changing rapidly in ways that call for new thinking by psychologists. The 13 articles in the special issue summarize current trends and knowledge and present new ideas for research, practice, and policy.

The issue covers a range of topics that explore how adult development is intertwined with cultural and historical change. These include how identity develops around reproductive and career concerns; the challenges of balancing the demands of work and family life; increases in stress associated with aging, caregiving, and economic issues; how changes in the workplace are reshaping the timing and experience of retirement; how digital technology is changing social relationships; and the importance of new positive narratives about aging.

What is the significance of this issue?

The course of adulthood has changed radically over recent decades. Pathways of education, work, and family life are more open and diverse than ever, and in some ways they are more stressful and challenging. Technology is reshaping how relationships and jobs change over the adult lifespan. Life expectancy is increasing, along with the potential for more healthy years following the exit from full-time work.

New theories and studies of adult development are needed to accommodate this increased diversity and unpredictability and to make sense of the societal shifts that have driven these changes.

Tell us about a few key takeaways.

There is greater diversity in the nature and pathways of adult development now than in the past. The special issue illustrates a multidisciplinary approach that considers factors such as culture, birth cohort, socioeconomic status, gender, race, and ethnicity to characterize and advance our understanding of adult development.

Although this makes it more complex and challenging to study the adult years, it also makes for a richer and more complete picture that can provide a useful framework for research and practice in the 21st century.

The global aging of societies calls for new perspectives and provides opportunities for addressing ageism, working longer, providing meaningful roles for older adults, and acknowledging the importance and ramifications of caregiving and grandparenting. At the same time there are challenges associated with living longer in the economic, physical health, mental health, and interpersonal spheres.

The special issue raises possibilities for new initiatives to highlight the range of circumstances and explore solutions.

What are some practical implications of the articles featured in the issue?

The articles in this special issue address distinctive challenges and opportunities faced by those in early, middle, and later adulthood. Psychosocial resources for dealing with vulnerabilities such as loneliness, economic loss, unemployment, loss or illness of loved ones, retirement, age discrimination, and aging-related declines are discussed.

The special issue considers how social disparities and stress are increasing and affecting mental and physical health. The articles address risk and resilience in the face of economic, physical, and mental health challenges.

Although the articles were written and accepted for publication before the COVID-19 pandemic, the content of the special issue is relevant for the post-COVID-19 world of adult development; these themes are likely to ring true as adults of all ages face many of these issues going forward.

Note: This article is in the Core of Psychology topic area. View more articles in the Core of Psychology topic area.

Arnett, J. J., Robinson, O., & Lachman, M. E. (2020). Rethinking adult development: Introduction to the special issue. American Psychologist, 75 (4), 425–430. https://doi.org/10.1037/amp0000633

About the Authors

Jeffrey Jensen Arnett is a senior research scholar at Clark University and executive director of the Society for the Study of Emerging Adulthood (SSEA). He has published widely on emerging adulthood as well as on the psychology of globalization and adolescent risk behavior.

Margie E. Lachman is the Minnie and Harold Fierman Professor of Psychology at Brandeis University. She is director of the Lifespan Development Lab and the Boston Roybal Center for Active Lifestyle Interventions. Her research focuses on optimizing physical, cognitive, and psychological health in middle and later adulthood.

Oliver C. Robinson is senior lecturer in psychology at the University of Greenwich, president of the European Society for Research in Adult Development, and author of Development through Adulthood . His research focuses on how aging, life transitions and crises affect identity, curiosity, wellbeing, and spirituality.

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Home Blog Everything You Needed to Know About Adult Learning Theory

Everything You Needed to Know About Adult Learning Theory

Research suggests that the adult brain starts getting lazy at around 25 years old . Of course, that doesn’t mean we aren’t capable of learning new concepts. It just means we need to engage in things that keep the brain busy and keep us learning.

Hobbies are one way to accomplish this. They can improve creative thinking, memory, and job performance. It’s easy for adults to revisit the learning process. We just need to rely on different strategies to retain the material.

In this blog, we unravel the intricacies, motivations, and strategies that shape how grown minds grasp new skills and insights. We’ll introduce the theory’s main pillars along with a few competing theories as they apply to adult learning needs.

Table of Contents

What Is Adult Learning Theory?

Adult Learning Theory is a framework that goes beyond the one-size-fits-all model. It delves into the diverse needs of adults, considering factors like prior experiences, responsibilities, and personal motivations. By acknowledging these nuances, educators can craft tailored strategies that make learning not only effective but also engaging and relevant.

American educator Malcolm Knowles introduced the concept back in 1968. Today, it is more commonly known as Adult Learning Theory.

The Importance of Adult Learning Theories

Unlike the traditional pedagogical approaches used with children and teenagers, adult learners bring a unique set of experiences, motivations, and challenges to the table. Adult Learning Theory takes center stage to decipher this enigma and provides educators and trainers with insights that can reshape education.

But why is this theory so vital? The answer lies in its power to bridge the gap between theory and practice. By understanding how adults learn best, educators can design courses, programs, and training sessions that tap into the real-world context of learners. This isn’t just about transferring knowledge; it’s about equipping adults with skills they can immediately apply to their careers, hobbies, and everyday lives.

The 8 Pillars of Adult Learning Theory

The principles of adult learning, often referred to as Andragogy, were formulated by Malcolm Knowles, a prominent figure in the field of adult education. These principles provide a framework for understanding how adults learn best. There are several adult learning principles, and the number varies depending on the source. Here are eight of the most commonly cited principles:

• Adults are self-directed learners.
• Adults have a wealth of experience that can be used to facilitate learning.
• Adults need to know why they are learning something and how it will help them achieve their goals.
• Adults learn best when the learning is relevant to their lives and work.
• Adults learn by doing and prefer hands-on experiences.
• Adults are ready to learn when they perceive a need for new knowledge or skills.
• Adults learn best when the learning is problem-centered and focused on real-world issues.
• Adults learn best when they are intrinsically motivated.

The Difference Between Adult Learning with Childhood Learning

Understanding the differences between how adults and children learn is essential for educators and trainers to design effective learning experiences for adult learners. Here’s an overview of the fundamental distinctions between these two learning processes:

Additional Adult Learning Theories

Click here to learn more about hands-on learning and how it might apply to business management.

Though Knowles’ Adult Learning Theory remains an incredibly influential method of instruction, competing theories do exist. These theories are important components of instructional design.

The more familiarity individuals responsible for our learning experiences have with them, the better they can instruct a diverse cast of adult learners.

Transformative Learning

Introduced in 1970 by sociologist and professor Jack Mezirow, this adult learning theory revolves around the premise that adults can adjust their thinking based on new information.

The theory, also known as “transformational learning,” is reserved for adults, as they possess the necessary set of experiences needed to undergo a significant thought transformation.

Mezirow says his theory revolves around two specific focuses: instrumental learning and communicative learning. The first relates to task-oriented problem-solving, while the second emphasizes how people communicate their feelings, needs, and desires.

Both elements are central to the transformation process, as students must embrace new perspectives that are both logical and emotional to challenge their previous understanding.

There is, however, a certain amount of criticism that follows Mezirow’s transformative learning theory. Concerns have been raised surrounding how the theory interacts with cultural contexts, relationships, and feelings. While new perspectives are often welcome, it’s difficult to accept them as a means of altering existing principles.

Self-Directed Learning

Self-directed learning ( SDL ) has roots in Andragogy. The theory maintains that adults must exercise control over learning decisions. Educators may act as supporting guides, but it’s up to the individual learner to take the initiative.

The idea here is that all students are responsible for their cognitive development. Students must be ready to self-regulate and self-evaluate their learning goals. This model is often used to describe eLearning environments.

Experiential Learning

Though he relied on existing theories to craft this adult learning theory, David Kolb introduced the concept of experiential learning back in 1970. This adult learning theory reflects a more hands-on approach, with actual experiences needed to make sense of new information.

In place of memorizing facts and statistics, adults will rely on real-world learning experiences to initiate reflection, review, and abstract thinking. They can then conclude and conceptualize the meaning of the experience.

Project-Based Learning

Project-Based Learning (PBL) was developed by American philosopher John Dewey back in 1897. This adult learning theory revolves around the concept of “learning by doing.”

Learners must demonstrate their knowledge by completing a project or overcoming an obstacle. The idea is that the adult learner will have an easier time retaining new information by completing a task than listening to someone explain it.

Find out what it means to attend a technical school today. Learn more about the kinds of learning styles they use.

Action Learning

Like many other theories on this list, action learning revolves around solving problems. Under this theory, learners need to exercise their critical thinking skills to isolate the problem and identify the solutions needed to resolve it. Only after these two steps have been completed should they take action.

This theory can facilitate learning in a group setting. This forces participants to both think critically and work collaboratively. Often, it is practiced in the workplace or across campuses.

Cooperative and Collaborative Learning

This theory applies to groups of two or more individuals sharing a common goal. It relies on collaborative skills, direct interaction, interdependence, personal accountability, and group interaction. Past experiences help learners form goals, resolve conflicts and post questions to the group.

Discovery Learning

This theory states adults learn best by formulating their own questions and answers. Originally introduced by Jerome Bruner, the theory encourages practitioners to rely on past experiences, knowledge, and even intuition.

Though instructors play a helpful role, it is up to the student to discover new information, correlations, and truths. Instead of absorbing recited information, learners should seek out original answers to their questions.

Elaboration Learning

Educational theorist Charles Reigeluth founded the elaboration theory. The model suggests that information should be presented in a specific order, from the most rudimentary to the most complex.

The idea is to help learners recognize connections between interrelated ideas. This method remains particularly popular among corporate learners.

Social Learning

Social learning theory was introduced back in the 1970s by psychologist Albert Bandura. The concept emphasizes the importance of observing, modeling, and imitating others’ behaviors and emotional reactions. The theory draws on both cognitive and behavioral elements.

Individualized Learning

This theory states that learners must be able to navigate topics independently to fully understand the ideas they involve. In a classroom setting, this typically involves some kind of assessment at the end of a chapter or unit. Social learning activities are peppered in to help broaden the learner’s understanding.

Behaviorism

Behaviorism frames all learning experiences as responses to external stimuli. Learners, in this case, can be considered “blank slates,” which may develop specific behaviors based on their interactions with the environment.

In this case, innate or inherited factors will have very little impact on a student’s existing knowledge base. The philosophy stems from the work of B.F. Skinner.

Cognitivism

This theory operates in contrast to behaviorism, asserting that individuals actively partake in learning. This kind of mental processing involves language, concept formation, and information processing. Cognitive theorists believe discrete changes in states of knowledge can measure learning.

Constructivism

Constructivism revolves around the idea that learners create meaning through experience. Though considered a branch of cognitivism, constructivism distinguishes itself through its unique definition of knowledge.

Constructivists believe we process new information through individual experiences. That means our catalog of knowledge is constantly subject to change.

Challenges of Adult Learning

Adult learners can face various challenges that can impact their learning journey. Recognizing and implementing strategies to address these challenges is crucial for creating effective adult education and training programs.

Here are some common challenges of adult learning, along with strategies that New England Tech offers to help adult learners overcome them:

Time Constraints

Adults often have busy schedules due to work, family responsibilities, and other commitments, which can make it difficult to allocate time for learning. Strategy: Offer flexible learning options, such as online courses or self-paced modules, that allow adult learners to fit learning into their schedules. Emphasize the value of small, consistent learning sessions.

Motivation and Relevance

Adult learners need to see the practical relevance of what they’re learning. If they can’t connect the content to their real-life situations, motivation can wane. Strategy: Clearly communicate the practical applications of the learning material. Incorporate case studies, real-world examples, and scenarios that resonate with their experiences.

Fear of Technology

Some adult learners may be uncomfortable with technology, particularly if they haven’t had much exposure to digital tools. Strategy: Provide user-friendly technology interfaces, offer tech support, and include tutorials to help learners navigate online platforms. Gradually introduce technology to build confidence.

Learning Styles and Preferences

Adults have diverse learning styles and preferences. A one-size-fits-all approach may not effectively engage all learners. Strategy: Offer a variety of learning materials and methods, including visual, auditory, and hands-on activities. Allow learners to choose the formats that suit them best.

Financial Constraints

Adult learners may have financial limitations that affect their ability to access learning resources or courses. Strategy: Offer affordable or accessible learning options whenever possible. Provide information about scholarships, grants, and financial assistance programs.

Learning Overload

Balancing learning with other responsibilities can lead to cognitive overload and hinder effective information retention. Strategy: Break down complex topics into smaller, manageable chunks. Use spaced repetition techniques to reinforce learning over time.

Limited Study Skills

Adult learners might not have well-developed study skills or effective learning strategies. Strategy: Incorporate study skills and learning techniques into the curriculum. Offer resources on time management, note-taking, and effective reading.

Adult Learning Theory isn’t just a concept; it’s a transformative force that has the potential to revolutionize the way we approach education for adults. It recognizes that the journey of learning doesn’t end with youth; instead, it evolves, adapts, and takes on new dimensions as we mature.

By acknowledging the autonomy of adult learners, valuing their prior experiences, and understanding their unique motivations, educators, and trainers can create learning experiences that resonate deeply. The principles of relevance, self-directed learning, and practical application become the pillars upon which effective education for adults is built.

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What exactly is Adult Learning Theory, and why is it essential for educators and trainers?

Adult Learning Theory is a framework that outlines how adults learn best, taking into account their unique experiences, motivations, and learning styles. It emphasizes principles like self-directed learning, relevance, and practical application. For educators and trainers, understanding this theory is essential because it enables them to create tailored, engaging, and effective learning experiences that resonate with adult learners, fostering better retention and application of knowledge in real-world contexts.

Is Adult Learning Theory applicable to all industries and professions?

Yes, Adult Learning Theory is applicable to all industries and professions. Its principles, such as recognizing prior experience, fostering relevance, and promoting self-directed learning, are universally relevant. Whether in corporate training, healthcare, technology, or any other field, understanding and implementing this theory enhances learning outcomes and engagement for adult learners across diverse contexts.

Can individuals benefit from understanding Adult Learning Theory for their personal development?

Absolutely, understanding Adult Learning Theory can greatly benefit individuals in their personal development. It provides insights into effective learning strategies, making self-guided learning more efficient. By recognizing one’s learning style, valuing prior experiences, and staying motivated through practical application, individuals can enhance their ability to acquire new skills, knowledge, and personal growth effectively.

How do you teach adults effectively?

Though different theories will emphasize different educational techniques, there are a few commonalities across the board. Most adult learning theories embrace the idea of angling lessons in ways that appeal to learners directly.

Drawing personal connections to information presented, keeping assignments relevant to the learners’ everyday responsibilities, and integrating existing academic training are good ways to approach adult education.

Why is Andragogy important?

Andragogy has been recognized as one of the first theories to distinguish adult learning patterns from how children digest new information. In other words, it was the first theory to point out that adults learn differently from kids.

This gives educators, coworkers, and even bosses much more direction when introducing new information to adult learners.

How do teachers use theories in teaching?

Teachers rely on adult learning theories when instructing older audiences in a variety of ways. These methods provide them with a basis to understand how their students learn.

At New England Tech, we know that everybody has their own way of learning. Our hands-on approach lets students learn and execute at the same time. Our faculty is prepared to work with students personally, armed with different tools to use when dealing with different kinds of learners.

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10 Simple Principles of Adult Learning

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Adult learning, or the act of pursuing one’s own education as an adult, can be done in a formal setting such as higher education, a trade school, or an apprenticeship. It can also refer to an adult simply wanting to learn about a new skill or topic independently. For most adults, pursuing education stems from a desire for self-improvement, a professional need for specific skills, or a want to expand available job possibilities.

When imagining the role of an educator, it’s easy to overlook adult education when you think about schools and learning. But adult education happens every day, and understanding the theory, challenges, and principles that go into adult learning is important.

What Is Adult Learning Theory?

In the 1980s, educator Malcolm Knowles popularized the concept of andragogy , the practice of teaching adults, and contrasted it with pedagogy, the practice of teaching children. The andragogy theory states that adult learners are vastly different from children in terms of their motivation, the relevancy of the education to their lives, and how they apply that education. In practice, adult learning focuses on giving adults an understanding of why they are doing something, lots of hands-on experiences, and less instruction so they can tackle things themselves. Many adult learning theories developed out of Knowles’ work in the following decades, all with the specific goal to enhance teaching methods and experiences for adult learners.

Why Are Adult Learning Theories Important?

For educators and educational institutions, being informed about the many challenges and theories behind effectively educating adults will only strengthen student outcomes and success. Children and adults are very different when it comes to how they learn, so different techniques must be used in order to make learning useful for adults.

And for adult learners getting ready to pursue higher education, knowing your learning style , understanding the strengths and weaknesses adult learners may have, and preparing for your individual strengths will also help you be successful. 

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Challenges of Adult Learning

Learning new skills or studying new subjects as an adult can be challenging for many reasons, most notably:

• Lack of time: Adults often have full-time jobs and sometimes children or other dependents relying on them. This can make finding the time to continue learning very difficult.
• Self-doubt: It’s common for adult learners to feel that they are too old to continue their education. They may feel it is too late and that they have missed their chance, but whether someone has five years or 50 years left in the workforce, they deserve to follow a passion and pursue a career that they’re excited about.
• Neuroplasticity: The human brain has an element of plasticity that helps one learn and grow. Younger people have brains that are more plastic, so changes are easier for them. As one ages, the brain becomes less plastic. That can result in a struggle for adult learners who are trying to take on new concepts, forge new pathways, and more. These learners may have a harder time understanding new things simply because their brains are less plastic. While this is a difficulty, it isn’t something that is insurmountable.
• Financial Barriers: Younger learners may have parental help when it comes to higher education. That’s usually not the case for adult learners. Finances can get in the way of learners pursuing their dream of earning a degree.
• Contradiction: Some of the things adult learners will learn in their education may be different than what they thought they knew or learned before. This can mean a shift in a person’s worldview, opinions, and knowledge, which requires mental flexibility.
• Lack of Support: It can be overwhelming to try and tackle earning a degree without a strong support system. Adult students may find they don’t have the emotional or social support they need in place to be able to tackle the difficulty of classes and learning.

What Are Adult Learning Principles? 

There are 10 simple principles of adult learning for future educators to keep in mind. All of these aspects are important when building curriculum and expectations for adult learners:

1. Adults Are Self-Directing:  For many adults, self-directed learning happens naturally without anyone explaining it or suggesting it. Adult learners are more prone to plan, carry out, and evaluate their learning experiences without the help of others. When instructing adults, it’s important for learners to set goals, determine their educational or training needs, and implement a plan to enhance their own learning.

2. Adults Learn by Doing : Many adults prefer not only to read or hear about subjects but to actively participate in projects and to take actions related to their learning. Project-based curriculum utilizes real-world scenarios and creates projects for students that they could encounter in a job in the future. Many adult learners find that this kind of learning is hugely beneficial for them as they apply what they have been taught to their careers, giving them direct access to seeing what they can do with their knowledge. 

3. Adults Desire Relevance: While some enjoy learning as an end in itself, adult learners are more likely to engage in learning that has direct relevance to their lives. For example, if they’re taking a certification course to improve their chances of promotion on the job, then the course should immediately address their needs. 

4. Adults Utilize Experience:   Adults are shaped by their experiences, and the best learning comes from making sense of those experiences. Adult learners can greatly benefit from finding ways to get hands-on learning. Internships, job shadowing opportunities, projects, and other experiential opportunities can help them get a firmer grasp of their learning and be more excited about how what they learn can be applied to their interests and careers.

5. Adults Process with Their Senses: Most adult learners don’t thrive as well in a lecture-style environment. Due to the lack of brain plasticity in older learners, it’s important to fully engage the senses when learning to successfully solidify new knowledge. Learning practices need to incorporate audio, visual, reading/writing, kinesthetic, independent, and group techniques. 

6. Adults Appreciate Repetition:  Repetition is essential for adult learning. If learners can practice new skills in a supportive environment, self-efficacy will develop to take those skills outside of the classroom. And the more they can practice a particular subject or skill, the better the chances are for mastery.

7. Adults Guide Their Own Development:   Utilizing dilemmas and situations to challenge an adult learner’s assumptions and principles helps them guide their own development. Adults can use critical thinking and questioning to evaluate their underlying beliefs and assumptions and learn from what they realize about themselves in the process. 

8. Adults Thrive with Goal Setting:  Learners who have a specific career or personal goal in mind will have a better experience as they pursue their degree programs. For example, if a student wants to learn Spanish before a trip to Mexico, they might have a specific goal to be conversational by a certain date. Adult learners need these goals because their learning is more in their own hands than younger learners.

9. Adults Learn Differently Than Children:  Children and adults are very different when it comes to how they learn, so different techniques must be used in order to make learning effective for adults. In addition to reading and memorizing, adult learners utilize their past life experiences and their current understanding of a subject as they learn. Also, adult learning needs to be problem-centered, making the impact more focused on current events or real life.

10. Adults Require Ownership:  With a more nuanced and advanced hierarchy of needs, adult learners place more value on intrinsic motivation and personal ownership of their learning. It's important to give adults internal motivation by recognizing their success and promoting increased self-esteem and confidence.

Ready to Learn More?

Do you want to impact the next generation of learners? Then check out WGU’s online teaching degree programs . Whether you’re looking to begin a career in education or to further your expertise, WGU’s teaching degrees provide the knowledge to step confidently into the classroom in any of the 50 states. 

Have you considered going back to school to gain the skills needed for a new career? Are you finally at a place to focus on your education? There are many great reasons to become an adult learner. With low, flat-rate tuition and a competency-based education model, WGU offers the support and flexibility needed to earn a degree at your own pace and comfort level.

Ready to Start Your Journey?

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Emerging Adult Essay: Life Moving Forward: A Personal Perspective on Transitioning Into Adulthood

• Published: May 2017
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Throughout life, transitions can create some of the most daunting experiences anyone will encounter. Leaving school, finding a job, moving out on your own, finding someone to share your life with—these all constitute life transitions and events that force everyone to look outside of their comfort zone to formulate a solution. Everyone dreads the change, but we all must face it head-on and accept the challenge of something different. For people with disabilities these changes may be overwhelming and, at times, debilitating. The key to success while transitioning is having attained the proper skill set to help overcome whatever transitional barrier(s) your particular disability mandates.

I am a 25-year-old male who was born and raised in Virginia. I am someone who, I hope and believe, has successfully transitioned to adulthood, though not without both ups and downs. Because I was born with cerebral palsy (CP), I use a power wheelchair to help me get around and accomplish daily tasks. I graduated from college in the fall of 2013. There I created my own disability advocacy degree under the interdisciplinary studies major titled “Advocacy for Social Justice.” I also majored in sociology and have a minor in psychology. Some of my hobbies include hanging out with friends and family, watching sports of all kinds, and working with individuals with disabilities. Until recently, when I opened my consulting business, I never held a full-time job, having only interned at all of my employment experiences. These internships include two federal and one state government entity and a disability resource center. Additionally, I have worked on a congressional campaign. I hope my experiences help to motivate you, whether you’re a student or a teacher reading this essay, to take control of your transition process or to help someone effectively transition to adulthood.

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Adult Learning, Essay Example

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The focus of this post is on adult learn ring as espoused by Knowles. According to the post Knowles advances that adult learning styles are different from those of children or adolescents. This is so because adults have a larger reservoir of experiences. It is very true. For example, if an adult learner is being taught computer skills the  learning techniques from other experiences will be applied in facilitating this activity receptivity.  The keys techniques from a piano key board may be applied in articulation of writing word or finding sites on the internet. Children or adolescents have a distinct application to the learning of computer skills and may even learn faster than an adult who brings a wealth of experiences into the activity (Larsen-Freeman, 2013). This post was written with prudence enlightening the reading audience.

Larsen-Freeman, D. (2013). Transfer of learning transformed. Language Learning: A Journal of Research in Language Studies, 63:S1

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The Relationship Between Emerging Adulthood as an Individual Mindset and Educational Outcomes: Current Issues and Directions for Future Research

• Original Article
• Published: 17 June 2024

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• Oxana Mikhaylova   ORCID: orcid.org/0000-0002-0236-6992 1  

This article reviews research on emerging adulthood (EA) and education to encourage scholars to further explore this topic. Specifically, this work focuses on EA as measured using the Inventory of the Dimensions of Emerging Adulthood (IDEA). Based on this review, it is concluded that the existing research provides mixed information regarding the link between education and EA mindset traits as measured by the IDEA. Furthermore, the connections between IDEA scores and both students’ academic achievements and the characteristics of the country’s educational system remain unclear. In addition, non-traditional forms of education have not been analyzed in terms of their association with IDEA scores. Finally, moderators and mediators of the relationship between EA as measured by the IDEA and education are worth examination in the future, and these could include gender, age, ethnicity, race, and social relationships.

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Mikhaylova, O. The Relationship Between Emerging Adulthood as an Individual Mindset and Educational Outcomes: Current Issues and Directions for Future Research. Trends in Psychol. (2024). https://doi.org/10.1007/s43076-024-00386-z

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Adult learning is of great interest to many researchers because continuous education is essential for people’s employment and social status. So, one should discuss the methods that help them acquire new knowledge. In particular, it is necessary to examine the environment in which learning can take place.

Adult learners can be viewed as a diverse group of people who differ in terms of their educational needs. For instance, these individuals represent various occupations. These employees have to extend their knowledge on a regular basis. Additionally, some companies encourage their workers to obtain graduate degrees. This task is critical for those people who want to assume the positions of leadership. Furthermore, these adults belong to various cultures. For instance, one can refer to immigrants who need to adjust to the new culture or language. Overall, they can prefer different learning environments. Some of them choose to attend formal institutions such as colleges or universes. In contrast, others seek the assistance of non-governmental organizations (NGOs) and community services. Moreover, they can pursue their studies without any external support.

There are three types of learning settings. At first, one should mention formal learning taking place in the institutions that are officially recognized. These organizations have a right to give certificates indicating that a person has some educational qualifications and skills (Merriam, Caffarella, & Baumgartner, 2007). In this case, the learning activities of a person follow the subject-based curriculum. Some types of formal education are compulsory. One should also consider the learning in non-formal settings. For instance, researchers mention the education offered by museums or libraries (Coffield, 2000, p. 12). Additionally, NGOs support people living in developing countries. For instance, they can help these individuals achieve computer literacy. These activities do not enable a person to receive the educational certificate that will be officially recognized. Moreover, the assessment standards are less rigorous. Finally, it is necessary to mention informal learning that accompanies the daily activities of a person (Sockett, 2014). These studies are usually self-directed. As a rule, a person does not have to follow the official curriculum. Furthermore, in many cases, there is no external assessment. Therefore, this learning does not lead to certification.

Overall, I would like to talk about informal learning that has been of great significance to me. For instance, I had to work very hard to get a good grasp of academic writing. In particular, I tried to study the essays of prominent writers and journalists. So, I attempted to emulate some of the techniques that they used. I was not required to do it, and my learning activities were not formally assessed. Nevertheless, these self-directed studies were of great use to me. In particular, I achieved higher scores for other papers that I submitted.

This discussion shows that adults can benefit from various learning environments. However, more attention should be paid to their motivation and willingness to acquire new skills. Overall, one should integrate these approaches to achieve the best results.

Reference List

Coffield, F. (2000). The Necessity of Informal Learning . New York, NY: Policy Press.

Merriam, S., Caffarella, R., & Baumgartner, L. (2007). Learning in Adulthood: A Comprehensive Guide . San Francisco, CA: Jossey-Bass.

Sockett, G. (2014). The Online Informal Learning of English . New York, NY: Palgrave Macmillan.

• Dorothea Orem's Philosophy of Self-Directed Care
• Self-Directed Learning Competence
• RL Wolfe Company's Self-Directed Work Teams
• Spirituality, Adult Education & Social Change
• American Association for Adult and Continuing Education
• Frustration and Resistance among Adult Learners
• Heutagogy and Lifelong Learning
• Adult Education and Lifelong Learning
• Chicago (A-D)
• Chicago (N-B)

IvyPanda. (2021, April 10). Adult Learning Environment. https://ivypanda.com/essays/adult-learning-environment/

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IvyPanda . (2021) 'Adult Learning Environment'. 10 April.

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1. IvyPanda . "Adult Learning Environment." April 10, 2021. https://ivypanda.com/essays/adult-learning-environment/.

Bibliography

IvyPanda . "Adult Learning Environment." April 10, 2021. https://ivypanda.com/essays/adult-learning-environment/.

• Research article
• Open access
• Published: 18 June 2024

Loss of protein tyrosine phosphatase receptor delta PTPRD increases the number of cortical neurons, impairs synaptic function and induces autistic-like behaviors in adult mice

• Bastián I. Cortés 1 ,
• Rodrigo C. Meza 2 ,
• Carlos Ancatén-González 2 , 3 ,
• Nicolás M. Ardiles 2 ,
• María-Ignacia Aránguiz 1 ,
• Hideaki Tomita 4 , 11 ,
• David R. Kaplan 4 , 5 ,
• Francisca Cornejo 6 ,
• Alexia Nunez-Parra 7 ,
• Pablo R. Moya 8 , 9 ,
• Andrés E. Chávez 2 , 10 &
• Gonzalo I. Cancino   ORCID: orcid.org/0000-0001-8697-7282 1  

Biological Research volume  57 , Article number:  40 ( 2024 ) Cite this article

Metrics details

The brain cortex is responsible for many higher-level cognitive functions. Disruptions during cortical development have long-lasting consequences on brain function and are associated with the etiology of brain disorders. We previously found that the protein tyrosine phosphatase receptor delta Ptprd, which is genetically associated with several human neurodevelopmental disorders, is essential to cortical brain development. Loss of Ptprd expression induced an aberrant increase of excitatory neurons in embryonic and neonatal mice by hyper-activating the pro-neurogenic receptors TrkB and PDGFRβ in neural precursor cells. However, whether these alterations have long-lasting consequences in adulthood remains unknown.

Here, we found that in Ptprd+/- or Ptprd-/- mice, the developmental increase of excitatory neurons persists through adulthood, affecting excitatory synaptic function in the medial prefrontal cortex. Likewise, heterozygosity or homozygosity for Ptprd also induced an increase of inhibitory cortical GABAergic neurons and impaired inhibitory synaptic transmission. Lastly, Ptprd+/- or Ptprd-/- mice displayed autistic-like behaviors and no learning and memory impairments or anxiety.

Conclusions

These results indicate that loss of Ptprd has long-lasting effects on cortical neuron number and synaptic function that may aberrantly impact ASD-like behaviors.

Introduction

The cerebral cortex is responsible for processing complex sensory and cognitive behaviors, requiring an extraordinary control over the generation of neurons and their connections during the development of the nervous system [ 1 ]. Typically, cortical circuits are assembled from different glutamatergic and GABAergic neuron lineages, each with unique morphological and physiological characteristics [ 2 , 3 ]. Indeed, disruption in the generation of neurons or their connectivity has long-lasting consequences affecting behavior that can result in brain disorders. In this regard, it has been proposed that the number, diversity, and function of glutamatergic and GABAergic neurons are essential to maintain the balance between different cerebral circuits and to avoid the onset of multiple neuropsychiatric conditions [ 4 ]. Perturbation of cellular processes such as neural precursor cell proliferation, neurogenesis, and neuronal migration during brain development may, in part, cause structural brain alterations that lead to the onset of brain disorders [ 5 ]. Gain or loss of expression or activity of genes involved in these processes are associated with the onset of several brain disorders, including autism spectrum disorder (ASD), schizophrenia, bipolar disorder, attention-deficit/hyperactivity disorder (ADHD), and intellectual disability [ 5 , 6 , 7 , 8 , 9 , 10 ]. We reported that alterations in the expression of one of these genes, protein tyrosine phosphatase receptor delta (PTPRD), impairs cerebral cortex development in mice [ 11 , 12 ].

Ptprd is a receptor protein tyrosine phosphatase type II family member. Structurally, it contains extracellular immunoglobulin and fibronectin III domains and two cytoplasmic tyrosine phosphatase domains [ 13 , 14 , 15 ]. We reported that Ptprd hetero- or homozygosity during cortical development triggered dysregulation of TrkB and PDGFRβ tyrosine kinase activity in embryonic and neonatal cortical neural precursor cells. Ptprd acted as a TrkB and PDGFRβ phosphatase, fine tuning kinase activity and preventing their hyperactivation. Loss of Ptprd resulted in aberrantly increased proliferation of neural precursor cells and concomitant mispositioning of their progeny, embryonic and newborn excitatory neurons [ 11 , 12 ]. Ptprd regulation of receptor tyrosine kinases like TrkB thus ensures the generation of appropriate numbers of neural precursor cells and neurons postnatally. Ptprd also performs a critical function at synapses, regulating cell adhesion important for synaptic formation and differentiation [ 15 ]. Given the role of Ptprd in assuring appropriate numbers of postnatal cortical neuron subtypes and its role at synapses, we asked if alterations of cortical neuron numbers observed embryonic and postnatally persist into adulthood, and if they perturb cortical synaptic function and ultimately, behavior.

Studies of how Ptprd deficiency alters excitatory and inhibitory synaptic transmission have yielded contradictory results [ 16 , 17 , 18 , 19 ]. For example, in hippocampal neuron cultures, Ptprd is involved in inhibitory synaptic formation and transmission but not in the development of excitatory synaptic transmission [ 16 ]. More recently, the loss of Ptprd expression differentially suppressed excitatory synaptic transmission in the stratum lacunosum moleculare in the hippocampus but not in the stratum radiatum [ 17 ]. Moreover, deletion of Ptprd in cultured neurons did not alter the development or number of synapses [ 18 ] and did not interfere with excitatory or inhibitory synaptic development and transmission [ 19 ]. It is important to note that these findings focused on hippocampal neurons. Whether the absence of PTPRD alters synaptic transmission in cortical neurons has not yet been addressed.

Here we address these issues by assessing in adult Ptprd-/- mice the number of glutamatergic and GABAergic neurons in the brain cortex, recording both excitatory and inhibitory synaptic function and analyzing animal behavior. Our results indicate that ablation of Ptprd results in an increase in both glutamatergic and GABAergic neurons in adult cortex leading to changes in excitatory and inhibitory synaptic transmission. Moreover, Ptprd+/- or Ptprd-/- mice show deficits in sociability and social novelty, and repetitive behavior characteristic of ASD-like behaviors when compared to Ptprd+/+ mice. These findings indicate that loss of one or both alleles of Ptprd during the formation of the cerebral cortex leads to profound structural changes that affect the assembly and functioning of glutamatergic and GABAergic circuits and synaptic transmission, likely contributing to ASD-like behaviors in adulthood.

The C57BL/6 N-A < tm1Brd >  Ptprd  < tm2a(KOMP)Wtsi>/WtsiOrl mice strain was purchased from Wellcome Trust Sanger Institute and maintained as heterozygous. The cross between heterozygous mice allowed the Ptprd+/+ , Ptprd+/- , and Ptprd-/- mice to be obtained. The following primers were used to identify the three genotypes mentioned above: Ptprd_111547_F: 5’-TCACCTCGCTGTTCTTCCTG-3’; Ptprd_111547_R:5’-CTTCTCAGTGCCCAACCCTC-3’; CAS_R1_Term: 5’-TCGTGGTATCGT TATGCGCC-3’. In addition, for sociability and social novelty tests, BALB/c wild-type mice were used as “old mouse” and “new mouse”. All mice had free access to rodent chow and water in a 12-hour dark-light cycle room with temperature controlled at 20–22 °C.

Perfusion of mice and brain extraction

Ptprd+/+ , Ptprd+/- , and Ptprd-/- mice were perfused transcardially at 3 months old. The mice were first placed in an isoflurane chamber to be lightly anesthetized. Once the animal was numb, the heart was exposed, and a saline solution (0.9% NaCl) was injected gradually through the right ventricle. Once the blood flow stopped, the brains were extracted and fixed in a 4% PFA solution overnight at 4 °C. The next day, the brains were dehydrated in 20%, 25%, and 30% sucrose (Merk, 107,651,100) solutions at 4 °C for 24 h each incubation. Subsequently, brains were embedded in the optimal cutting temperature compound (OCT; Sakura, 4583) and stored at -80 °C until cut. Next, 18 μm coronal slices were obtained by cryostat (Leica, CM1850) at a temperature of -20 °C and stored at -80 °C until use.

Immunofluorescence and quantification

For morphometric analysis, immunostaining of tissue sections was performed as described [ 11 , 12 , 20 ]. Briefly, brain sections were washed with 1X TBS for immunostaining. Then were permeabilized with TBS − 0.3% Triton X-100 for 30 min. The tissues were then incubated in TBS with 5% BSA 0.3% Triton X-100 for 1 h as a blocking solution. Brain slices were incubated overnight with primary antibodies in a blocking solution at 4 °C. The next day, the sections were washed with TBS and then incubated with secondary antibodies in a blocking solution for 1 h at room temperature. After TBS washes, sections were counterstained with Hoechst 33,258 for 10 min and mounted. To evaluate the number of GABAergic neurons in the somatosensory cortex and medial prefrontal cortex (mPFC), rabbit anti-parvalbumin antibody (1:100; Swant, PV 25) and mouse anti-somatostatin antibody (1:500; Santa Cruz, sc-55,565) were used. To evaluate the number of glutamatergic neurons in the somatosensory cortex and mPFC, rabbit anti-Tbr1 antibody (1:400; Abcam, ab31940) and rabbit anti-Satb2 antibody (1:500; Abcam, ab92446) were used. To quantify the specific neuronal types, the somatosensory cortex and mPFC boundaries were established using the mouse brain Atlas [ 21 ] and Adobe Photoshop CC 2015. Then, the images were quantified using the Cell Counter tool of the Fiji program.

Prefrontal cortex slice preparation

Acute coronal brain slices (300 μm thick) were obtained as previously described [ 22 ]. Briefly, the brain was removed and slices containing the mPFC were cut using a DTK-1000 Microslicer (Ted Pella, Inc.) in an ice-cold high-choline solution (110 mM Choline-Cl, 2.5 mM KCl, 1.25 mM NaH 2 PO 4 , 7 mM MgCl 2 , 25 mM NaHCO 3 , 15 mM glucose, 0.5 mM CaCl 2 , 11.6 mM ascorbate, 3.1 mM pyruvate (290–305 mmol/Kg)). Thirty minutes post-sectioning, coronal slices were gradually switched to artificial cerebral spinal fluid (aCSF) solution (124 mM NaCl, 2.69 mM KCl, 1.25 mM KH 2 PO 4 , 1.3 mM MgSO 4 , 26 mM NaHCO 3 , 10 mM glucose, 2.5 mM CaCl 2 , pH 7.4 (300–305 mmol/Kg)). All solutions were equilibrated with 95% O 2 and 5% CO 2 (pH 7.4), and slices were kept at room temperature for at least 30 min before recording.

Electrophysiology

All experiments, unless otherwise stated, were performed at 28 ± 1 °C in a submersion-type recording chamber perfused at ~ 2 mL/min with aCSF supplemented with either the GABAA receptor antagonist picrotoxin (PTX; 50 µM) or the AMPA/Kainate and NMDA receptor antagonists CNQX (25 µM) and D-APV (50 µM) to block fast inhibitory and excitatory transmission, respectively. Coronal slices were visualized using infrared differential interference contrast on a Nikon Eclipse FN1 microscope, and layer 2/3 pyramidal neurons from prelimbic (PrL) and infralimbic (IL) subdivisions of mPFC were morphologically identified. Whole-cell voltage-clamp recordings using a Multiclamp 700B amplifier (Molecular Devices, Sunnyvale, CA, USA) were made from 2/3 pyramidal neuron somas located ~ 200 mm from pia and voltage-clamped at -60 or 0 mV (unless otherwise stated) using patch-type pipette electrodes (~ 3.0–4.5 MΩ) containing 131 mM Cs-gluconate, 8 mM NaCl, 1 mM CaCl 2 , 10 mM EGTA, 10 mM glucose, 10 mM HEPES, 5 mM MgATP, and 0.4 mM Na 3 GTP; pH 7.2–7.4 (285 mmol/kg). To assess cell stability, series, and input resistances were monitored with test pulses (-4mV, 80 ms) throughout all experiments. Cells with > 20% change in series resistance were excluded from the analysis.

To elicit excitatory and inhibitory postsynaptic currents (EPSCs and IPSCs, respectively), a bipolar-stimulating patch pipette filled with aCSF was placed in layer 2/3 around 100–150 μm from the recording neuron and voltage pulses (1–15 V, 100–200 ms pulse width) were delivered through a stimulus isolator (DS2A-Mk.II, Digitimer Ltd). Typically, stimulation was adjusted to obtain comparable magnitude synaptic response across experiments (e.g., 100–150 pA EPSC and IPSC). Two different protocols were used to evaluate short-term synaptic plasticity: First, two pulses at different interstimulus intervals (10, 30, 70, 100, and 300 ms) were used to calculate the paired-pulse ratio (PPR) that was defined as the ratio of the amplitude of the second response to the amplitude of the first response. Second, synaptic depression was evaluated using a burst of 25 (for EPSC) and 20 (for IPSC) stimuli at 14 and 10 Hz, respectively, and delivered every 60 s; 5 burst-evoked responses were averaged for each experiment. Miniature EPSCs (mEPSCs) were recorded at 32 ± 1 °C from neurons voltage-clamped at -60 mV in the continuous presence of tetrodotoxin (TTX, 500 nM). In contrast, isolated miniature IPSCs were recorded at 0 mV in the continuous presence of TTX, CNQX (25 µM), and D-APV (50 µM). mEPSCs and mIPSC were identified using a minimal threshold amplitude (≥ 5 pA) and analyzed using the mini-analysis software Synaptosoft (Synaptosoft). Spontaneous excitatory and inhibitory currents (sEPSCs/sIPSCs) were recorded without TTX, whereas AMPAR/NMDAR ratios were analyzed by recording AMPAR-mediated EPSCs at -60 mV. In contrast, NMDAR-mediated EPSCs were recorded at + 40mV in the continuous presence of CNQX (25 µM) to isolate NMDA-mediated EPSC. All drugs were obtained from Sigma and Tocris, prepared in stock solutions in milli-Q water or DMSO, and added to the ACSF as needed. Total DMSO in the aCSF was maintained at > 0.1%. Synaptic EPSCs and IPSCs were elicited at 10-sec intervals (0.16 Hz), filtered at 2.2 kHz, and acquired at 10 kHz using custom-made software written in Igor Pro 4.09 A (WaveMetrics).

Morris water maze test

Morris water maze navigation task was performed to evaluate spatial learning and memory as previously described [ 23 ]. Briefly, in the learning phase, the animals were trained to locate a hidden 9-cm-diameter white platform (1 cm below the surface of the water) for 4 consecutive days following 4 external cues, 4 times per day per animal in a 1.2-m-diameter circular pool filled with 23 ± 2 °C painted white water. Each trail ended when the animals found the platform and remained there for 10 s. If the animal did not find the platform at the end of 60 s, they were gently guided there for the remaining 10 s. Later in each trial, the animals were gently removed from the maze, dried with a tissue paper towel, and put in a similar housing cage with additional tissue paper towels to continue drying. On the fifth day, in the memory phase, the test was performed by removing the platform and allowing the free swimming for 60 s. The behavior was monitored during all tasks using an automatic tracking system (ANY-maze video tracking software, Stoelting Co, Wood Dale, IL, USA). The learning phase measured latency and path length to the platform, while the memory test measured the time spent in each quadrant and the target quadrant and the traveled distance within the target quadrant.

Y-Maze test

Memory was also evaluated with the Y-maze as previously described [ 24 ]. Briefly, Y-Maze tests were performed using a maze with 3 arms of 8 cm width x 35 cm long and walls of 20 cm height, connected at a 120° angle, shaping a Y and assigned as arms A, B, and C. Each animal was placed in the arm (A), looking to the center of the maze, and tracked for 8 min by the ANY-maze software. Four clues were put on the maze walls to guide the animals. Alternation was defined as a visit to 3 different arms (ABC, ACB, BCA, BAC, CAB, CBA), and % spontaneous alternation was determined by the following formula:

% Spontaneous alternation = (Total alternations / (Total arm entries-2) x 100.

Open field test

Anxiety was evaluated with the open field test as described previously [ 25 ]. Briefly, mice were exposed to a free exploration in an open field chamber of 40 cm (l) x 40 cm (w) x 40 cm (h). This arena is divided into two areas by a line: the peripheral and central regions. The time in each chamber area was recorded for 10 min by ANY-maze software.

Elevated plus maze test

Anxiety was also evaluated with the evaluated plus maze as described previously [ 26 ]. Briefly, the test was performed using a maze with 4 arms of 7.5 cm width x 35 cm long, shaping a cross with a 7.5 × 7.5 cm center and 50 cm from the floor. 2 arms (closed arms) had walls of 20 cm height, and the others (open arms) were uncovered. The animals were placed in the center of the maze and tracked for 5 min by ANY-maze software.

Self-grooming test

Repetitive behavior was measured by the self-grooming test as previously described [ 25 , 27 ]. Briefly, mice were recorded for 10 min through a 20 × 20 cm transparent box to allow visualization of the animal. The cumulative time spent by the animals performing the self - grooming process was quantified.

Marble burying test

Repetitive behavior was also evaluated by the marble burying test as previously described [ 22 , 25 ]. Briefly, the animals were exposed to a free exploration arena of 23 × 33 cm in the presence of 20 proportionally distributed black marbles. The marbles were placed in a 5 cm thick layer of dry sawdust to allow the animal to perform the digging behavior. After 30 min, the number of buried marbles was quantified. A marble was considered buried if 2/3 were covered.

Three-chamber test

The three-chamber social novelty and sociability test was used to evaluate deficits in social behavior as previously described [ 25 , 27 , 28 ]. First, the animals were subjected to 10 min of habituation in the three chambers. Then, to assess the sociability of the animals, mice were placed in a context in which they chose to interact with either a caged mouse (old mouse) or an inanimate object, each in a different chamber for 10 min. Subsequently, to assess the social novelty, the inanimate object was replaced by a second unfamiliar mouse (new mouse), and the experimental mouse was placed in a context in which it had to choose between interacting with a familiar or an unfamiliar mouse for 10 min. ANY-maze software recorded the time the experimental animals spent with each option.

Statistical analyses

Statistical analyses were performed using the one-way ANOVA test with unpaired data to compare data between Ptprd +/+, Ptprd +/-, and Ptprd -/- genotypes. Unless otherwise indicated, all electrophysiological results were statistically analyzed using paired and unpaired two-tailed Student’s t-test and One- or Two-way ANOVA at the p  < 0.05 significance level in OriginPro 7.0 and 8.6 (OriginLab). In the figures illustrated, traces are averages of 25–30 responses. In the case of the statistical analysis used to evaluate the sociability and social novelty test, the 2-way ANOVA test was used. Corrections for multiple comparisons were performed using Tukey’s post hoc test. All statistical tests were performed with Prism 6 software. In all cases, the bars indicate the standard error of the mean.

Ptprd +/- or Ptprd -/-mice display an increase in glutamatergic neurons and impaired synaptic function in the adult cortex

We previously reported that Ptprd+/- and Ptprd-/- embryos have more excitatory cortical neurons [ 11 ]. To determine whether this phenotype persisted until adulthood, we performed immunostaining for Tbr1 (Fig.  1 A) and Satb2 (Fig.  1 C), markers of deep- and upper-layer excitatory cortical neurons, respectively, in coronal sections of the mPFC, a brain area involved in different cognitive processes, from 3-month-old Ptprd+/+ , Ptprd+/- , and Ptprd-/- mice. While heterozygous mice showed no differences as compared to control littermates, Ptprd-/- mice had 20% more Tbr1-positive neurons than control and Ptprd+/+ littermates in the mPFC (Fig.  1 B). However, we did not observe any changes in the number of Satb2-positive neurons (Fig.  1 D), suggesting that only early born neurons are affected at the mPFC. To complement this result, we analyzed the somatosensory cortex, another cortical region commonly disrupted in brain disorders associated with PTPRD mutations. We found that Ptprd-/- and Ptprd+/- mice had more Tbr1-(Fig.  1 E, F) and Satb2-positive excitatory neurons (Fig.  1 G, H).

Adult  Ptprd+/- or Ptprd-/-  mice show an increase of glutamatergic neurons in the mPFC and somatosensory cortex. ( A-H ) Coronal sections of mPFC ( A, C ) or somatosensory cortex ( E, G ) of Ptprd +/+ (PTPRD-WT) and Ptprd -/- (PTPRD-KO) 3-month-old animals were immunostained against Tbr1 ( A, E ) or Satb2 ( C, G ). Relative total Tbr1-positive cells in the mPFC ( for B , PTPRD-WT, n  = 3; PTPRD-HET, n  = 4; PTPRD-KO, n  = 4 ) or somatosensory cortex ( for F , PTPRD-WT, n  = 3; PTPRD-HET, n  = 5; PTPRD-KO, n  = 5 ) of Ptprd +/+, Ptprd +/- (PTPRD-HET), and Ptprd -/- 3-month-old mice were quantified. Relative total Satb2-positive cells in the mPFC ( for D , PTPRD-WT, n  = 3; PTPRD-HET, n  = 3; PTPRD-KO, n  = 4 ) or somatosensory cortex ( for H , PTPRD-WT, n  = 3; PTPRD-HET, n  = 4; PTPRD-KO, n  = 5 ) of Ptprd +/+, Ptprd +/-, and Ptprd -/- 3-month-old mice were quantified. In ( A ) and ( C ), the dotted line shows mPFC and specific cortical layers. (I, J) Tbr1 ( I ) or Satb2 ( J ) positive neurons were quantified per cortical layers II/III, V and VI in the mPFC. (K, L) Tbr1 ( K ) or Satb2 ( L ) positive neurons were quantified per cortical layers II/III, IV, V and VI in the somatosensory cortex. mPFC = medial prefrontal cortex. SSC = somatosensory cortex. WM = white matter. * p  < 0.05; ** p  < 0.01; *** p  < 0.001. In all the images, scale bars represent 200 μm. In all graphs, the error bars denote SEMs

The brain cortex has 6 layers which are connected to different brain circuits. To determine if this increase in excitatory neurons is cortical layer specific [ 1 , 3 , 29 ], the mPFC and somatosensory cortex were divided into their respective layers as previously described [ 21 , 30 , 31 , 32 , 33 ]. In the mPFC, there were no differences in the number of Tbr1- or Satb2-positive excitatory neurons per cortical layer (Fig.  1 I, J). However, in the somatosensory cortex, the increase in Tbr1-positive neurons was specific to layers IV and VI in 3-month-old Ptprd+/- or Ptprd-/- mice compared to Ptprd+/+ mice (Fig.  1 K). For Satb2-positive neurons, the increase was observed in all cortical layers in the somatosensory cortex of 3-month-old Ptprd-/- mice compared to Ptprd+/+ mice, except for cortical layer V, where Ptprd+/- mice had more Satb2-positive neurons compared to Ptprd+/+ mice (Fig.  1 L). These results indicate that loss of Ptprd in adult mice results in an increase in excitatory cortical neurons in brain areas associated with cognition.

As an increased number of glutamatergic neurons is associated with altered synaptic function [ 33 ], we characterized the impact of the Ptprd ablation on the spontaneous synaptic activity in layer 2/3 pyramidal neurons in the mPFC. Firstly, we recorded spontaneous excitatory synaptic currents (sEPSCs) and found a significant increase in the frequency but not amplitude of sEPSCs in Ptprd-/- and not Ptprd+/- mice when compared to Ptprd+/+ littermates (Fig.  2 A). Similarly, an increase in the frequency of miniature excitatory synaptic currents (mEPSCs) but not in the amplitude was observed in Ptprd+/- or Ptprd-/- mice compared to Ptprd+/+ mice (Fig.  2 B). Secondly, we analyzed the evoked synaptic transmission mediated by AMPA receptors by generating input–output curves. We observed an increased response at high voltage inputs in Ptprd-/- mice compared to Ptprd+/+ (Fig.  2 C), suggesting that excitatory transmission is augmented in the Ptprd-/- mice. Thirdly, we examined whether the effect of the PTPRD ablation alters synaptic release probability by estimating two principal forms of short-term synaptic plasticity: paired-pulse ratio (PPR) and use-dependent depression. However, both PPR (Fig.  2 D) and synaptic depression in response to a stimulus train of 25 stimuli at 14 Hz (Fig.  2 E) remained unchanged between genotypes. Finally, we measured the AMPA/NMDA ratio and found no significant differences between genotypes (Fig.  2 F). Together, these results suggest that the increase in excitatory transmission cannot be accounted for by changes in postsynaptic receptor numbers or changes in release probability, but rather, could be due to an increase in the number of synaptic contacts in Ptprd-/- mice.

Excitatory synaptic transmission in layer 2/3 of the mPFC is disrupted in the Ptprd-/-  mice. ( A ) Sample traces (left) and quantitative analysis (right) of sEPSC activity recorded from layer 2/3 pyramidal neurons in the mPFC in Ptprd +/+ (PTPRD-WT), Ptprd+/- (PTPRD-HET), and Ptprd -/- (PTPRD-KO) mice. A significant increase in the frequency but not in the amplitude of sEPSC in PTPRD-KO was detected. PTPRD-WT, n  = 4 animals, 10 cells; PTPRD-HET, n  = 5 animals, 11 cells; PTPRD-KO, n  = 8 animals, 15 cells. ** p  < 0.01. ( B ) Miniature excitatory synaptic current (mEPSC) also shows an increase in the frequency but not in the amplitude in Ptprd-/- compared to Ptprd+/+ and Ptprd+/- mice. PTPRD-WT, n  = 5 animals, 8 cells; PTPRD-HET, n  = 5 animals, 8 cells; PTPRD-KO, n  = 6 animals, 11 cells. * p  < 0.05. ( C ) Input/output function measured as the EPSP amplitude as a function of stimulus intensity also is increased in Ptprd-/- compared to Ptprd+/+ and Ptprd+/- mice. PTPRD-WT, n  = 3 animals, 6 cells; PTPRD-HET, n  = 5 animals, 9 cells; PTPRD-KO, n  = 5 animals, 7 cells. ( D ) Paired-pulse responses superimposed after subtraction of the first pulse at 10, 30, 70, 100, and 300 ms interstimulus intervals (ISI) are similar between genotyping. PTPRD-WT, n  = 4 animals, 6 cells; PTPRD-HET, n  = 6 animals, 8 cells; PTPRD-KO, n  = 5 animals, 8 cells. ( E ) Short-term synaptic responses (top) and normalized summary data (bottom) evoked by a burst of 25 stimuli at 14 Hz are also similar between genotyping. PTPRD-WT, n  = 3 animals, 6 cells; PTPRD-HET, n  = 4 animals, 8 cells; PTPRD-KO, n  = 4 animals, 9 cells. ( F ) AMPA/NMDA ratio shows no significant differences between genotyping. PTPRD-WT, n  = 3 animals, 6 cells; PTPRD-HET, n  = 3 animals, 8 cells; PTPRD-KO, n  = 5 animals, 14 cells. In all graphs, the error bars denote SEMs

Ptprd +/- or Ptprd -/- mice display an increase in GABAergic neurons and aberrant inhibitory synaptic transmission in the adult cortex

Ptprd is expressed in subcortical structures during brain development [ 34 ] and is important for GABAergic synaptic differentiation [ 35 , 36 , 37 ]. However, its role in regulating the generation of GABAergic neurons has not been described. We therefore evaluated whether loss of Ptprd expression could also alter the number of GABAergic neurons in the adult mouse cortex. To address this, we performed immunostaining for parvalbumin (Fig.  3 A) and somatostatin (Fig.  3 C), two major types of cortical GABAergic neurons [ 2 ], at the mPFC. We found that Ptprd-/- mice had more parvalbumin-positive (Fig.  3 B) and somatostatin-positive neurons (Fig.  3 D) than wild-type littermates in the mPFC. Additionally, we immunostained for parvalbumin (Fig.  3 E) and somatostatin (Fig.  3 G) at the somatosensory cortex. Quantification showed that Ptprd-/- mice had more parvalbumin- (Fig.  3 F) and somatostatin-positive neurons (Fig.  3 H). In addition, Ptprd heterozygous mice had more somatostatin-positive neurons (Fig.  3 H), indicating that GABAergic neurons are increased in the cortex of 3-month-old Ptprd-/- mice.

Ptprd+/- or Ptprd-/- mice show an increase of GABAergic neurons in the mPFC and somatosensory cortex at 3 months. ( A-H ) Coronal sections of the mPFC or somatosensory cortex of Ptprd +/+ (PTPRD-WT) and Ptprd -/- (PTPRD-KO) 3-month-old animals were immunostained against parvalbumin ( A, E ) or somatostatin ( C, G ). Relative total parvalbumin-positive cells in the mPFC ( for B , PTPRD-WT, n  = 4; PTPRD-HET, n  = 3; PTPRD-KO, n  = 5 ) or somatosensory cortex ( for F , PTPRD-WT, n  = 4; PTPRD-HET, n  = 5; PTPRD-KO, n  = 3 ) of Ptprd +/+, Ptprd +/- (PTPRD-HET), and Ptprd -/- 3-month-old mice. Relative total of somatostatin-positive cells (arrows head) in the mPFC ( for D , PTPRD-WT, n  = 4; PTPRD-HET, n  = 3; PTPRD-KO, n  = 4 ) or somatosensory cortex ( for H , PTPRD-WT, n  = 3; PTPRD-HET, n  = 4; PTPRD-KO, n  = 4 ) of Ptprd +/+, Ptprd +/- and Ptprd -/- 3-month-old mice. In ( A ) and ( C ), the dotted line shows mPFC and specific cortical layers. (I, J) Parvalbumin ( I ) or Somatostatin ( J ) positive neurons were quantified per cortical layers II/III, V and VI in the mPFC. (K, L) Parvalbumin ( K ) or Somatostatin ( L ) positive neurons were quantified per cortical layers II/III, IV, V and VI in the somatosensory cortex. mPFC = medial prefrontal cortex. SSC = somatosensory cortex. PV = parvalbumin. SOM = somatostatin. WM = white matter. In all the images, scale bars represent 200 μm, and arrows show positive cells. * p  < 0.05; ** p  < 0.01; *** p  < 0.001. In all graphs, the error bars denote SEMs

We then evaluated the number of inhibitory neurons per layer. We found no differences in the number of parvalbumin-positive neurons in the mPFC per layer across genotypes (Fig.  3 I). However, when we analyzed the somatostatin-positive neurons, we found that they were increased specifically in the mPFC layer V of 3-month-old Ptprd+/- or Ptprd-/- compared to Ptprd+/+ mice (Fig.  3 J). Next, we performed a similar analysis in the somatosensory cortex and found that parvalbumin-positive neurons were specifically increased in layers V and VI (Fig.  3 K). For somatostatin-positive neurons, the increase was observed in all cortical layers in the somatosensory cortex of 3-month-old Ptprd-/- compared to Ptprd+/+ mice, except for cortical layers V and VI, where Ptprd+/- mice also had more somatostatin-positive neurons compared to Ptprd+/+ mice (Fig.  3 L).

As the increase in glutamatergic neurons was coupled with increase synaptic activity, we evaluated whether the increase in GABAergic neurons observed in Ptprd-/- mice was associated with an increase in inhibitory synaptic transmission by recording spontaneous inhibitory synaptic currents. As we found with excitatory synaptic transmission (Fig.  2 ), we observed an increase in the frequency but not amplitude of spontaneous inhibitory synaptic currents (sIPSCs) in Ptprd-/- but not Ptprd+/- mice when compared to Ptprd+/+ littermates (Fig.  4 A). Similarly, there was an increase in the frequency but not amplitude of miniature synaptic currents (mIPSCs) in Ptprd-/- mice (Fig.  4 B). Next, we evaluated evoked inhibitory input-output curves and found an enhancement in evoked IPSCs in Ptprd-/- compared to Ptprd+/+ mice (Fig.  4 C), suggesting an increase in GABAergic synaptic transmission. To further evaluate the effect of the PTPRD ablation on GABAergic synaptic function, we measured PPR and use-dependent depression and found that both PPR and synaptic depression in response to a stimulus train of 20 stimuli at 10 Hz remained unchanged between genotypes (Fig.  4 D, E), indicating that the increase in GABAergic function is independent of changes in release probability.

Inhibitory synaptic transmission in layer 2/3 of the mPFC is impaired in the Ptprd-/-  mice. ( A ) Representative traces (left) and quantitative analysis (right) of sIPSC activity recorded from layer 2/3 pyramidal neurons in the mPFC in Ptprd +/+ (PTPRD-WT), Ptprd+/- (PTPRD-HET), and Ptprd -/- (PTPRD-KO). PTPRD-WT, n  = 5 animals, 9 cells; PTPRD-HET, n  = 6 animals, 8 cells; PTPRD-KO, n  = 4 animals, 9 cells. * p  < 0.05. ( B ) Representative traces (left) and quantitative analysis (right) of mIPSC activity recorded from layer 2/3 pyramidal neurons in the mPFC in Ptprd +/+ (PTPRD-WT), Ptprd+/- (PTPRD-HET), and Ptprd -/- (PTPRD-KO). PTPRD-WT, n  = 4 animals, 9 cells; PTPRD-HET, n  = 4 animals, 8 cells; PTPRD-KO, n  = 4 animals, 8 cells. * p  < 0.05. ( C ) Evoked IPSC amplitudes as a function of stimulus intensity plotted as input/output curves show an increase in Ptprd-/- compared to Ptprd+/+ and Ptprd+/- mice. Significant changes between Ptprd+/+ and Ptprd-/- groups are as “*”. * p  < 0.05, ** p  < 0.01, and *** p  < 0.001. PTPRD-WT, n  = 4 animals, 7 cells; PTPRD-HET, n  = 5 animals, 9 cells; PTPRD-KO, n  = 4 animals, 8 cells. ( D ) Paired-pulse responses at 10, 30, 70, 100, and 300 ms interstimulus intervals (ISI) are similar between genotyping. PTPRD-WT, n  = 4 animals, 6 cells; PTPRD-HET, n  = 4 animals, 6 cells; PTPRD-KO, n  = 5 animals, 8 cells. ( E ) Short-term synaptic responses (top) and normalized summary data (bottom) evoked by a burst of 20 stimuli at 10 Hz are also similar between genotyping. PTPRD-WT, n  = 4 animals, 7 cells; PTPRD-HET, n  = 6 animals, 10 cells; PTPRD-KO, n  = 4 animals, 7 cells. ( F ) Excitatory and inhibitory balances show no significant differences between genotyping. PTPRD-WT, n  = 4 animals, 8 cells; PTPRD-HET, n  = 4 animals, 10 cells; PTPRD-KO, n  = 5 animals, 12 cells. In all graphs, the error bars denote SEMs

Finally, we tested whether the enhanced excitatory and inhibitory inputs onto layer 2/3 pyramidal neurons in Ptprd-/- mice altered the excitatory/inhibitory balance of these neurons. We recorded simultaneously evoked EPSC and IPSCs by holding pyramidal neurons at − 35 mV and found no significant differences in the excitation/inhibitory ratio among genotypes (Fig.  4 F), indicating that the increase in synaptic function observed in Ptprd-/- mice does not alter the excitatory/inhibitory balance in the mPFC.

Ptprd +/- or Ptprd -/- mice do not show impairments in learning and memory or increased anxiety

Mutation or aberrant expression of genes associated with neuroanatomical alterations are thought to have profound consequences in adulthood as they affect the connectivity between neurons, promoting brain disorders [ 5 ]. As PTPRD mutations have been associated with brain disorders that include cognitive impairments, we evaluated whether Ptprd+/- or Ptprd-/- mice have learning and memory deficits by performing the Morris Water Maze test. We found that Ptprd-/- or Ptprd+/- exhibited normal latency escape (Fig.  5 A). Also, they showed normal time spent (Fig.  5 B) and distance (Fig.  5 C) in the target quadrant as well as in the time spent (Fig.  5 D) and distance (Fig.  5 E) in the area where the platform was located compared to Ptprd+/+ mice . To confirm these observations, we performed the Y-maze test, and found that Ptprd+/- or Ptprd-/- mice showed no difference in the percentage of spontaneous alternation preference as compared to Ptprd +/+ animals (Fig.  5 F). These results indicate that Ptprd+/- or Ptprd-/- mice do not present with spatial learning and memory deficits.

Heterozygous or homozygous Ptprd  mice showed no learning and memory deficits. ( A-E ) 3-month-old Ptprd+/+ (PTPRD-WT), Ptprd +/- (PTPRD-HET), and Ptprd-/- (PTPRD-KO) mice were tested in the Morris water maze (MWM) test. (A) Latency escape during the 4 training days. (B) Time spent by the animals swimming in the target quadrant area on test day (day 5). (C) Distance traveled in the target quadrant zone on test day. (D) Time spent by the animals swimming in the platform area on test day. (E) Distance traveled in the platform zone on test day. PTPRD-WT, n  = 8; PTPRD-HET, n  = 16; PTPRD-KO, n  = 9. (F) 3-month-old Ptprd+/+ , Ptprd+/- , and Ptprd-/- mice were tested in the Y-maze test. Percentage of preference for spontaneous alternations (SAP). PTPRD-WT, n  = 6; PTPRD-HET, n  = 11; PTPRD-KO, n  = 8. In all graphs, the error bars denote SEMs

As individuals with PTPRD variants exhibit anxiety [ 38 ] we evaluated if Ptprd+/- or Ptprd-/- mice showed increased anxiety using the open-field and elevated plus maze tests [ 26 , 39 ]. In the open field test, we observed no differences between Ptprd+/- or Ptprd-/- and Ptprd +/+ animals in the time spent in the central zone (Fig.  6 A) and the number of times that mice entered this area (Fig.  6 B). In addition, none of the animal groups presented differences in the distance covered, indicating that the mice did not have motor deficits (Fig.  6 C). In the elevated plus maze test, there were no differences between genotypes in terms of the percentage of time spent in the open (Fig.  6 D) and closed arms (Fig.  6 E) or in the number of entries to these arms (Fig.  6 F, G). Together, these results suggest that losing one or both alleles of Ptprd does not result in learning and memory impairment or aberrant anxious behavior.

Loss of one or both Ptprd  alleles does not induce anxious behavior. ( A-C ) 3-month-old Ptprd+/+ (PTPRD-WT), Ptprd+/- (PTPRD-HET), and Ptprd-/- (PTPRD-KO) mice were tested in open-field anxiety test. (A) Percentage of the total time that the mice spent in the central zone. ( B ) Number of times that the animals enter the central zone. ( C ) Total distance traveled. PTPRD-WT, n  = 10; PTPRD-HET, n  = 16; PTPRD-KO, n  = 11. ( D-G ) 3-month-old Ptprd +/+, Ptprd +/-, and Ptprd -/- mice were tested in the elevated plus maze test (EPM). ( D ) Percentage of the total time spent in open arms. ( E ) Percentage of the total time spent in closed arms. (F) Number of times that the animals enter the open arms. ( G ) Number of times that the animals enter the closed arms. PTPRD-WT, n  = 10; PTPRD-HET, n  = 12; PTPRD-KO, n  = 11. In all graphs, the error bars denote SEMs

Ptprd +/- or Ptprd -/- mice exhibit autistic-like behaviors

PTPRD variants have also been associated with ASD [ 40 , 41 , 42 , 43 ]. Therefore we asked whether the loss of one or both Ptprd-/- alleles induces autistic-like behaviors. To address this, we evaluated social behavior using the three-chamber test for social interaction and novelty [ 44 ]. In this test, wild-type rodents prefer to spend more time with another rodent (sociability) and will investigate a novel intruder more than a familiar one (social novelty). To evaluate the sociability of Ptprd-/- mice, they were placed in a context where they could interact with a caged mouse or an inanimate object, each in a different chamber. We observed that Ptprd-/- or Ptprd+/- mice, unlike Ptprd +/+ animals, did not show a preference to spend more time exploring the chamber containing the caged mouse, indicating a social deficit (Fig.  7 A). The inanimate object was then replaced by another mouse (new mouse) and the caged mouse was kept in the same chamber (old mouse). Ptprd-/- or Ptprd+/- mice did not discriminate between the old and new mouse, indicating a deficit in social novelty compared to Ptprd+/+ mice (Fig.  7 B).

Ptprd+/- or Ptprd-/- mice showed autistic-like behaviors. ( A-D ) 3-month-old Ptprd +/+ (PTPRD-WT), Ptprd +/- (PTPRD-HET), and Ptprd -/- (PTPRD-KO) mice were tested for sociability ( A ) and social novelty ( B ), self-grooming ( C ) or marble burying ( D ). ( A ) Social Interaction Test was measured in the three-chamber test. The graph shows the cumulative duration of Ptprd +/+, Ptprd +/-, and Ptprd -/- mice in the mouse or object chambers. PTPRD-WT, n  = 12; PTPRD-HET, n  = 13; PTPRD-KO, n  = 9. ( B ) Social Novelty Test was assessed in the three-chamber test. The graph shows the cumulative duration of Ptprd +/+, Ptprd +/-, and Ptprd -/- mice in the old mouse and new mouse chambers. PTPRD-WT, n  = 12; PTPRD-HET, n  = 13; PTPRD-KO, n  = 9. * p  < 0.05; ** p  < 0.01. (C, D) Repetitive behavior was evaluated by the time spent in spontaneous self-grooming (C) or marble-burying behaviors ( D ). ( C ) Cumulative time in which Ptprd +/+, Ptprd +/-, and Ptprd -/- mice did self-grooming behavior in a total of 10 min. PTPRD-WT, n  = 7; PTPRD-HET, n  = 7; PTPRD-KO, n  = 8. ( D ) Percentage of marbles buried by Ptprd +/+, Ptprd +/-, and Ptprd -/- mice after 30 min. PTPRD-WT, n  = 7; PTPRD-HET, n  = 14; PTPRD-KO, n  = 7. * p  < 0.05; ** p  < 0.01. In all graphs, the error bars denote SEMs

Next, we evaluated repetitive and stereotyped behaviors. Self-grooming behavior is often altered in mouse models of ASD [ 45 ]. Thus, we recorded mice from below through a transparent box to allow the visualization and further analysis of self-grooming. Ptprd -/- mice increased their time self-grooming as compared to Ptprd+/+ mice (Fig.  7 C). Finally, we used the marble burying test, a test widely used to assess repetitive behavior in mouse models of compulsive disorder and ASD [ 46 , 47 ]. The test assesses the percentage of marbles that mice bury in 30 min, which indirectly reflects the degree of repetitive digging behavior. We found that Ptprd-/- or Ptprd+/- mice showed a significant increase in buried marbles as compared to Ptprd+/+ mice (Fig.  7 D). Therefore, our behavioral results suggest that loss of one or both Ptprd alleles induces autistic-like behaviors.

Here, we analyzed how loss of Ptprd expression impacts the generation of glutamatergic and GABAergic neurons and synaptic transmission in the cerebral cortex, potentially contributing to the emergence of behavioral impairments in adulthood. Mutations or aberrant expression of this gene have previously been associated with various neurodevelopmental disorders such as obsessive-compulsive disorder, schizophrenia, and ASD [ 14 ]. In this regard, we reported that Ptprd is a key regulator of mouse embryonic neurogenesis, controlling the numbers of neurogenic intermediate progenitor cells and, ultimately, the number of neonatal cortical neurons [ 11 , 12 ]. Here, we found that the developmental increase of excitatory neurons persisted through adulthood, affecting excitatory synaptic function in the mPFC. Moreover, loss of Ptprd expression resulted in an increase in inhibitory GABAergic neurons and inhibitory synaptic function. Finally, adult Ptprd-/- mice displayed autistic-like behaviors, suggesting that ablation of Ptprd not only influences the number and function of cortical neurons but also behavior.

What accounts for the increase in glutamatergic neurons in adult Ptprd-/- mice? We propose that it is directly related to the increase in Tbr2-positive intermediate precursor cells in these mice during embryogenesis that we previously reported [ 11 , 12 ]. There is abundant evidence that ASD-associated genes exhibit cell cycle changes in neural precursor cells during brain development, culminating in an alteration in the total number of glutamatergic neurons [ 48 ]. For example, Fang et al. [ 33 ], reported that the increase in Tbr2-positive intermediate precursors that in turn produces increased amounts of glutamatergic neurons, was sufficient to elicit alterations in synaptic function, interneuron mislocalization, and the appearance of ASD-like behaviors. The genes affected in animal models of ASD in which the number of Tbr2-positive cells is increased are many [ 48 ]. Therefore, given that the pathophysiological mechanisms that trigger the onset of ASD-like behaviors include alterations in the production of glutamatergic neurons [ 33 , 48 ] like those observed in the mPFC and somatosensory cortex of Ptprd+/- or Ptprd-/- mice, this could, at least in part, explain the ASD-like behavior in these animals.

Ptprd participates both as an adhesion molecule promoting synaptic formation and differentiation, and a phosphatase regulating the activity of receptors involved in synaptic transmission such as TrkB [ 14 ]. The role of Ptprd in synaptic function in cultured hippocampal neurons is conflicting, as the absence of Ptprd has been reported to either not alter synaptic formation or transmission [ 18 , 19 ] or decrease excitatory synaptic transmission [ 17 ]. Our results obtained in acute mPFC slices from adult Ptprd -/- mice indicate an increase in both excitatory and inhibitory synaptic activity. While this increase was not due to changes in the number of postsynaptic receptors or release probability, it is possible that these differences could be due to increased number of synaptic contacts in the Ptprd-/- mPFC.

Ptprd+/- or Ptprd-/- mice showed autistic-like behaviors. ASD is characterized by social deficits and repetitive behavior [ 49 ]. Consistent with this idea, Ptprd+/- or Ptprd-/- mice present deficits in sociability with a conspecific and the preference for interaction with a new individual, supporting findings of mutations in PTPRD linked to ASD risk [ 40 , 41 , 42 , 43 ]. Moreover, Ptprd-/- mice have increased self-grooming and marble-burying behavior, core symptoms of ASD. However, our findings do differ from those reported by Ho et al. [ 50 ] that did not report changes in the time invested in self-grooming. These differences likely reflect the different experimental paradigms used. We evaluated instinctively self-grooming behavior without stimuli to promote it. Ho et al. [ 50 ] quantified self-grooming time after spraying the animals with a sucrose solution to encourage the indicated behavior. We propose that artificially stimulating self-grooming could mask normal behavior in Ptprd+/+ animals. We also evaluated the number of the marbles buried that represents the occurrence of repetitive behavior, while Ho et al. [ 50 ] evaluated the time and number of times the experimental animals spent performing the digging process. While the experimental paradigms used by both groups are valid, our findings of increased repetitive behavior in the Ptprd-/- mice do indeed indicate that they exhibit ASD-like behavior.

Uetani et al. [ 51 ] and Drgonova et al. [ 52 ] reported impairments of spatial learning and memory in Ptprd-/- mice which we did not observe. While the Uetani et al. [ 51 ], Drgonova et al. [ 52 ], and our mice were in the same C57BL/6J genetic background, there were evident differences in the mouse models. The mice of those groups showed growth retardation and weight loss attributed to problems in food intake. While their Ptprd-/- mice lived for approximately one year, our animals aged normally. To confirm the lack of spatial learning and memory impairments, we also evaluated our animals in the Y-maze test, which showed no cognitive deficits associated with learning and memory. Our findings regarding the lack of anxious behavior in Ptprd-/- mice were similar to the results reported by Park et al. [ 17 ] and Ho et al. [ 50 ], who used the same Ptprd -/- model as that by Uetani et al. [ 51 ] and Drgonova et al. [ 52 ]. The findings will need to be resolved in future studies with neural precursor specific knockouts of Ptprd .

Our data, together with previous findings [ 11 , 12 ], show that several phenotypic features normally associated with ASD were present in mice lacking Ptprd , including deficits in neural precursor cell proliferation, increased neurogenesis, neuronal mislocalization [ 48 , 53 ], alterations in the number and function of glutamatergic and GABAergic neurons [ 54 ], and social deficits and repetitive behavior [ 55 ]. These findings lead us to propose that the absence of Ptprd expression disrupts the generation and functioning of glutamatergic and GABAergic cortical neurons, inducing the appearance of ASD-like behavior in adulthood in mice. Our results suggest a possible explanation for why variants in PTPRD result in brain disorders.

Data availability

The datasets used and/or analyzed during the current study are available from the corresponding author on reasonable request.

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Acknowledgements

We thank the technical support of Jessica Molina, Sebastián Bondi, María Beatriz Lazen, Ale Frene, Gloria Lucero, and Luis Lopez.

This work was supported by the research grants ANID-FONDECYT Regular 1201848 (to AEC), 1210507 (to GIC), and 1231012 (to PRM); ANID-FONDECYT Iniciación 11220708 (to FC); ANID-FONDECYT postdoctorado grant 3190793 (to RCM); CIHR to DRK, and ANID-FONDEQUIP (EQM160154 to AEC). CAG was supported by a Ph.D. fellowship from ANID 21201603.

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Bastián I. Cortés, María-Ignacia Aránguiz & Gonzalo I. Cancino

Centro Interdisciplinario de Neurociencia de Valparaíso (CINV), Facultad de Ciencias, Universidad de Valparaíso, Valparaíso, 2340000, Chile

Rodrigo C. Meza, Carlos Ancatén-González, Nicolás M. Ardiles & Andrés E. Chávez

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Francisca Cornejo

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Centro de Estudios Traslacionales en Estrés y Salud Mental (C-ESTRES), Universidad de Valparaíso, Valparaíso, 2340000, Chile

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Instituto de Fisiología, Facultad de Ciencias, Universidad de Valparaíso, Valparaíso, 2340000, Chile

Instituto de Neurociencias, Facultad de Ciencias, Universidad de Valparaíso, Valparaíso, 2340000, Chile

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Contributions

BIC, MIA, GIC and FC performed the neuroanatomical studies and their analyzes. RCM, AEC and CAG performed the electrophysiological experiments and their analyses. PRM, GIC, BIC, NA and HT performed the behavioral experiments and their analyzes. ANP, AEC, PRM, DRK and GIC conceptualized all the experiments, analyzed the data, and wrote the manuscript. All authors participated in the writing of the manuscript. All authors read and approved the final manuscript.

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Correspondence to Gonzalo I. Cancino .

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Animal use was approved by Universidad Mayor, Universidad de Valparaiso, and Pontificia Universidad Católica de Chile Bioethics and Animal Care Committees in accordance with the bioethics regulation of the Chilean Research Council (ANID).

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Cortés, B.I., Meza, R.C., Ancatén-González, C. et al. Loss of protein tyrosine phosphatase receptor delta PTPRD increases the number of cortical neurons, impairs synaptic function and induces autistic-like behaviors in adult mice. Biol Res 57 , 40 (2024). https://doi.org/10.1186/s40659-024-00522-0

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Enhancing the Learning Experience at Essex County College with Moodle Integration

This essay discusses the integration of Moodle at Essex County College, highlighting its benefits for enhancing the educational experience. Moodle, an open-source learning management system, fosters collaboration, accessibility, and personalized learning. It supports multimedia content, offers diverse assessment tools, and provides valuable analytics for educators. The platform’s user-friendly interface and robust security measures make it an effective tool for modern education, aligning with the college’s commitment to excellence and innovation.

How it works

Essex County College has long been a beacon of accessible education, serving a diverse student body with a commitment to excellence and innovation. As technology continues to reshape the educational landscape, the integration of Moodle, a powerful and versatile learning management system (LMS), offers a unique opportunity to enhance the learning experience for both students and educators at Essex County College.

Moodle, short for Modular Object-Oriented Dynamic Learning Environment, is an open-source platform designed to provide educators, administrators, and learners with a single robust, secure, and integrated system to create personalized learning environments.

The integration of Moodle at Essex County College can transform traditional classroom dynamics, making education more interactive, engaging, and accessible.

One of the most significant advantages of Moodle is its ability to foster a collaborative learning environment. Traditional classrooms often rely heavily on face-to-face interactions, which, while valuable, can be limiting in scope and scale. Moodle expands these interactions into the digital realm, offering forums, chat rooms, and collaborative wikis where students can engage with peers and instructors outside of scheduled class times. This extended interaction fosters a deeper understanding of course material, as students can continue discussions, ask questions, and work together on projects regardless of their physical location.

Moreover, Moodle’s integration can significantly enhance the accessibility of educational resources. In the past, students who missed a class due to illness or other commitments might struggle to catch up. With Moodle, all course materials, including lecture notes, reading assignments, and recorded lectures, can be uploaded and accessed anytime. This flexibility ensures that all students have the opportunity to stay current with their studies, promoting equity in education.

The platform’s support for multimedia content is another notable feature that can enrich the learning experience. Educators at Essex County College can incorporate videos, podcasts, and interactive simulations into their courses, catering to various learning styles and making complex concepts easier to understand. For instance, a biology professor could supplement a lecture on cellular processes with a high-resolution video of cell division, providing students with a visual and auditory learning aid that reinforces their understanding.

Assessment and feedback mechanisms within Moodle are designed to enhance the learning process. Traditional paper-based assessments can be time-consuming to grade and may not provide timely feedback. Moodle offers a variety of assessment tools, including quizzes, assignments, and peer assessments, that can be graded automatically. This immediate feedback loop helps students identify areas of improvement and allows instructors to tailor their teaching strategies to meet the needs of their students more effectively.

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Additionally, Moodle’s open-source nature means that it can be continuously improved and customized to meet the specific needs of Essex County College. The platform’s large global community of developers and users contributes to its ongoing development, ensuring that it remains at the cutting edge of educational technology. This collaborative approach aligns with the college’s commitment to innovation and continuous improvement.

Security and privacy are paramount considerations in the digital age, and Moodle’s robust security measures help protect sensitive student and faculty data. The platform adheres to stringent data protection standards, ensuring that personal information is kept secure. This level of security builds trust among users and supports a safe online learning environment.

In conclusion, the integration of Moodle at Essex County College represents a significant step forward in enhancing the learning experience for students and educators alike. By fostering collaboration, increasing accessibility, supporting diverse learning styles, and providing valuable insights into student performance, Moodle creates a dynamic and inclusive educational environment. The platform’s user-friendly interface, personalized learning paths, and robust security measures further contribute to its effectiveness as a learning management system. As Essex County College continues to embrace technological advancements, Moodle stands as a powerful tool that supports the institution’s mission of providing high-quality education and fostering student success.

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PapersOwl.com. (2024). Enhancing the Learning Experience at Essex County College with Moodle Integration . [Online]. Available at: https://papersowl.com/examples/enhancing-the-learning-experience-at-essex-county-college-with-moodle-integration/ [Accessed: 18-Jun-2024]

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Surgeon General: Why I’m Calling for a Warning Label on Social Media Platforms

By Vivek H. Murthy

Dr. Murthy is the surgeon general.

One of the most important lessons I learned in medical school was that in an emergency, you don’t have the luxury to wait for perfect information. You assess the available facts, you use your best judgment, and you act quickly.

The mental health crisis among young people is an emergency — and social media has emerged as an important contributor. Adolescents who spend more than three hours a day on social media face double the risk of anxiety and depression symptoms, and the average daily use in this age group, as of the summer of 2023, was 4.8 hours . Additionally, nearly half of adolescents say social media makes them feel worse about their bodies.

It is time to require a surgeon general’s warning label on social media platforms, stating that social media is associated with significant mental health harms for adolescents. A surgeon general’s warning label, which requires congressional action, would regularly remind parents and adolescents that social media has not been proved safe. Evidence from tobacco studies show that warning labels can increase awareness and change behavior. When asked if a warning from the surgeon general would prompt them to limit or monitor their children’s social media use, 76 percent of people in one recent survey of Latino parents said yes.

To be clear, a warning label would not, on its own, make social media safe for young people. The advisory I issued a year ago about social media and young people’s mental health included specific recommendations for policymakers, platforms and the public to make social media safer for kids. Such measures, which already have strong bipartisan support, remain the priority.

Legislation from Congress should shield young people from online harassment, abuse and exploitation and from exposure to extreme violence and sexual content that too often appears in algorithm-driven feeds. The measures should prevent platforms from collecting sensitive data from children and should restrict the use of features like push notifications, autoplay and infinite scroll, which prey on developing brains and contribute to excessive use.

Additionally, companies must be required to share all of their data on health effects with independent scientists and the public — currently they do not — and allow independent safety audits. While the platforms claim they are making their products safer, Americans need more than words. We need proof.

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