Students in their first semester at university were asked to pose their own questions, known as quecture questions, for each learning topic. This was as part of their weekly preparation for flipped lectures on two parallel compulsory biology courses. Quecture questions are intended to engage students, particularly those with educational disadvantages who may be unfamiliar with effective learning strategies, in meaningful concept-based learning. The mechanism of posting the questions differed between the two courses in that a forced sequence tool on the virtual learning environment required students to post something on the online board on one course while students could opt not to post on the online board on the other course. Small focus groups and qualitative analysis showed that the students realised both cognitive and metacognitive learning benefits from the quecture strategy. Students that were first-in-family to Higher Education were prioritised as focus group attendees and, although adjustment to the quecture strategy was judged to be more difficult within this group, positive learning benefits were discerned for all these students. The pressure to engage within one course was, on balance, considered to be useful for nudging students towards more effortful engagement with their learning. In this way cognitive and metacognitive benefits were recognised for first-in-family students.

Background Quectures are flipped lectures with embedded 'quecture questions', which employ metacognitive reflection as an active learning intervention. Quecture questions are students' own questions, identified and constructed by students around learning objectives during lectures. The quecture question intervention aims to support each student to engage better with their learning and to work at an individually appropriate level, and is accessible to all students irrespective of learning background. This research explores engagement and performance of students using the intervention, with a focus on those who traditionally receive low scores, to measure the effectiveness of quecture questions. Results Accumulated data demonstrate that student engagement with quecture questions is associated with improved learning on wider topics than those on which the intervention was used, as evidenced by improved overall course scores. Students who participate minimally within other elements of the course but who do submit their own quecture questions gain disproportionate benefit. The improved performance benefits of engagement with quecture questions are most marked for students with low prior scores. Students originating from the same country as our university and those whose parents did not attend university represent typically low-scoring demographic groups with low participation levels who might potentially benefit from the quecture strategy. However, lack of engagement with the intervention is also prevalent in these student groups precluding unengaged students from benefitting. Conclusions Quecture questions represent a simple yet effective intervention for improving learning in lectures. Broad learning benefits suggest improved learning habits, and support the notion that the intervention leads to students assuming increased responsibility for their own learning. The use of quecture questions generated most benefit to low-scoring students, validating the intervention as a useful tool with which to address learning inequities. The quecture question intervention readily adapts to suit online learning and represents an ideal first step for busy instructors wishing to adapt their lectures towards a more student-centred approach to learning.

The transition into higher education stretches students socially, academically and within their personal lives requiring adaptation and development of resilience. For many, such demands may lead to decreased mental well-being and, for some, mental ill-health.This project aimed to trial simple mental health awareness and well-being tasks with first year undergraduate students, and to determine whether students find these interventions beneficial and worthy of embedding as transition activities within the first-year curriculum. Four activities were trialled with 185 first-year students who reported the activities as beneficial. All activities caused an overall increase in student knowledge of how to maintain good mental well-being. In the light of this project’s findings, such activities are recommended for embedding into the first-year curriculum and throughout higher education.

“Graduate attributes” are widely believed to be important in developing the scientific skill-set, with generic skills being viewed as more important than discipline-specific qualities. Importantly, students need opportunities to think and practice in ways akin to experts. The continued use of didactic lectures in university education often leads to the accumulation of superficial knowledge, and does not adequately train students to acquire the skills and attributes required of an effective scientist: critical thinking, an inquiring mind and creativity. We analysed active learning lecture strategies in a second year genetics course to determine their effectiveness in developing the scientific skill-set. These were found to be more beneficial than standard lecturing. Investigation of one of these strategies, the “quecture” (an adaptation of the flipped classroom), found that students did not view this method as being the most useful, despite being the most interactive. Our evidence suggests this student resistance to result from the requirement for prior preparation, perceived as an increased workload. We advocate the incorporation of active learning strategies in lectures to support the development of students’ scientific skill–set and specifically advise the introduction of novel formats such as the quecture early in university level science education. Keywords: Active learning, interactive engagement, quecture, scientific skill-set, graduate attributes

Active learning exercises engage students during lectures, but often fail to take account of the individual learning position of each student. The ‘quecture’ is a partially flipped lecture that incorporates students posing their own questions (quecture questions), discussing them during lectures and revisiting them later. These interactive learning events are designed to personalise students’ construction of learning during lectures. Quectures were trialled in direct comparison with both fully flipped and traditional lectures, providing information on student attitudes, experiences and engagement with the learning strategy. Quectures were favoured by participants over the two other lecture formats and were found to be helpful both in increasing learning and in improving study habits, although some students had difficulty adjusting to, or disliked, the new mode of learning. The student-posed questions were also perceived by students to improve enquiry skills and to personalise learning. Although many chose not to engage with the strategy, those who did felt more engaged with, and more responsible for their own learning during quectures than in traditional lectures. Future work will be required to generalise the effectiveness of this strategy as well as to fine tune for optimum benefit. It will also be important to investigate which subpopulations of students preferentially engage or disengage with the strategy, and to unpick any relationship between this engagement and academic performance.

BACKGROUND AND RATIONALE Closed (student only) Facebook groups are commonplace amongst students and the social and academic benefits of such groups are well studied. The exclusion of staff from these Facebook groups could conceivably lead to inappropriate behaviour such as plagiarism but the occurrence of this is unknown. Many students have a poor understanding of what constitutes plagiarism, or the rationale for its avoidance (other than to avoid penalty). In this study we sought to explore the behaviours of students on Facebook groups, focusing on plagiarism. We also broadened our analysis to encompass an investigation of general plagiarism awareness in order to use the findings to inform the co-creation of a simple plagiarism intervention tool for use on Facebook groups. METHODS Our mixed methods approach encompassed seven student focus groups and a survey of 273 students at one UK University, as well as consultation of 11 HEI staff. Information and ideas drawn from the investigative phase of the project informed the design and co-creation of a plagiarism avoidance video resource. RESULTS AND LESSONS LEARNED We did not find plagiarism on Facebook to be a major concern despite wide use by students of closed groups to support learning. We did, however, find that interactive discussion on Facebook was minimal, and that membership was non-inclusive, and therefore suggest caution towards formal use of Facebook in learning. We also found that >40% of our students held misconceptions and anxieties about what constitutes plagiarism and about how to avoid it, and that University guidelines on plagiarism are not improving understanding. Plagiarism confusion was heightened for students new to our University, although further research into the generality of this observation is required. We advocate good scholarship education that actively engages, but is not limited to, students in their first year of study, and present our own anti-plagiarism tool that is suitable for deployment via Facebook groups.

General rights Copyright for the publications made accessible via the Edinburgh Research Explorer is retained by the author(s) and / or other copyright owners and it is a condition of accessing these publications that users recognise and abide by the legal requirements associated with these rights. Take down policy The University of Edinburgh has made every reasonable effort to ensure that Edinburgh Research Explorer content complies with UK legislation. If you believe that the public display of this file breaches copyright please contact openaccess@ed.ac.uk providing details, and we will remove access to the work immediately and investigate your claim.

We demonstrate that student engagement with PeerWise, an online tool that allows students to author and answer multiple‐choice questions (MCQs), is associated with enhanced academic performance across diverse assessment types on a second year Genetics course. Benefits were consistent over three course deliveries, with differential benefits bestowed on groups of different prior ability. A rating scheme, to assess the educational quality of students' questions, is presented and demonstrates that our students are able intuitively to make such quality assessments, and that the process of authoring high quality questions alone does not explain the academic benefits. We further test the benefits of providing additional PeerWise support and conclude that PeerWise works efficiently with minimal intervention, and can be reliably assessed using automatically generated PeerWise scores. © 2014 by The International Union of Biochemistry and Molecular Biology, 42(5):371–381, 2014.

The relationship between students' use of PeerWise, an online tool that facilitates peer learning through student-generated content in the form of multiple-choice questions (MCQs), and achievement, as measured by their performance in the end-of-module examinations, was investigated in 5 large early-years science modules (in physics, chemistry and biology) across 3 research-intensive UK universities. A complex pattern was observed in terms of which type of activity (writing, answering or commenting on questions) was most beneficial for students; however, there was some evidence that students of lower intermediate ability may have gained particular benefit. In all modules, a modest but statistically significant positive correlation was found between students' PeerWise activity and their examination performance, after taking prior ability into account. This suggests that engaging with the production and discussion of student-generated content in the form of MCQs can support student learning in a way that is not critically dependent on course, institution, instructor or student.

We describe one aspect of a UK inter-institutional project wherein an online tool was used to support student generation of multiple choice questions. Across three universities and in five modules in physics, chemistry and biology, we introduced the PeerWise online system as a summative assessment tool in our classes, the desire being to increase student engagement, academic attainment and level of cognitive challenge. Engagement with the system was high with many students exceeding the minimum requirements set out in the assessment criteria. We explore the nature of student engagement and describe a working model to enable high-impact student-learning and academic gain with minimal instructor intervention.

In the mammalian model of sex determination, embryos are considered to be sexually indifferent until the transient action of a sex-determining gene initiates gonadal differentiation. Although this model is thought to apply to all vertebrates, this has yet to be established. Here we have examined three lateral gynandromorph chickens (a rare, naturally occurring phenomenon in which one side of the animal appears male and the other female) to investigate the sex-determining mechanism in birds. These studies demonstrated that gynandromorph birds are genuine male: female chimaeras, and indicated that male and female avian somatic cells may have an inherent sex identity. To test this hypothesis, we transplanted presumptive mesoderm between embryos of reciprocal sexes to generate embryos containing male: female chimaeric gonads. In contrast to the outcome for mammalian mixed-sex chimaeras, in chicken mixed-sex chimaeras the donor cells were excluded from the functional structures of the host gonad. In an example where female tissue was transplanted into a male host, donor cells contributing to the developing testis retained a female identity and expressed a marker of female function. Our study demonstrates that avian somatic cells possess an inherent sex identity and that, in birds, sexual differentiation is substantively cell autonomous.

In 2001 it was established that, contrary to our previous understanding, a mechanism exists that equalises the expression levels of Z chromosome genes found in male (ZZ) and female (ZW) birds (McQueen et al. 2001). More recent large scale studies have revealed that avian dosage compensation is not a chromosome-wide phenomenon and that the degree of dosage compensation can vary between genes (Itoh et al. 2007; Ellegren et al. 2007). Although, surprisingly, dosage compensation has recently been described as absent in birds (Mank and Ellegren 2009b), this interpretation is not supported by the accumulated evidence, which indicates that a significant proportion of Z chromosome genes show robust dosage compensation and that a particular cluster of such dosage compensated genes can be found on the short arm of the Z chromosome. The implications of this new picture of avian dosage compensation for avian sex determination are discussed, along with a possible mechanism of avian dosage compensation.

Birds undergo genetic sex determination using a ZW sex chromosome system. Although the avian mechanisms of neither sex determination nor dosage compensation are understood, a female-specific non-coding RNA (MHM) is expressed soon after fertilisation from the single Z chicken chromosome and is likely to have a role in one or both processes. We have now discovered a prominent female-specific modification to the Z chromatin in the region of the MHM locus. We find that chicken chromatin at Zp21, including the MHM locus, is strongly enriched for acetylation of histone H4 at lysine residue 16 in female but not male chromosomes. Interestingly, this specific histone modification is also enriched along the length of the up-regulated Drosophila melanogaster male X chromosome where it plays a vital role in the dosage compensation process.

The genes MBD1 and MBD2 encode methyl-CpG binding proteins that suppress transcription from methylated promoters. In contrast, CGBP encodes a protein that binds promoters containing unmethylated CpG and stimulates transcription. All three are located on human chromosome 18q21, a region of frequent loss of heterozygosity in several cancers. These genes therefore represent candidate tumour suppressor genes, whose loss of function could affect the normal regulation of gene expression, whether by lack of complete suppression of genes normally silenced (via loss of MBD1 and MBD2 ) or by some loss of activation of genes normally expressed (via loss of CGBP ), either way contributing to the tumorigenic phenotype. We have confirmed by fluorescent in situ hybridization that MBD1 and MBD2 bracket the DCC locus giving a gene order of MBD1 / CGBP–DCC 5′- DCC 3′- MBD2 . Mutation analyses by single-stranded conformation polymorphism in colon and lung cancer cell lines and primary tumours revealed a small number of mutations, suggesting only a limited role of these genes in human tumorigenesis.

The Z and W sex chromosomes of birds have evolved independently from the mammalian X and Y chromosomes [1]. Unlike mammals, female birds are heterogametic (ZW), while males are homogametic (ZZ). Therefore male birds, like female mammals, carry a double dose of sex-linked genes relative to the other sex. Other animals with nonhomologous sex chromosomes possess "dosage compensation" systems to equalize the expression of sex-linked genes. Dosage compensation occurs in animals as diverse as mammals, insects, and nematodes, although the mechanisms involved differ profoundly [2]. In birds, however, it is widely accepted that dosage compensation does not occur [3-5], and the differential expression of Z-linked genes has been suggested to underlie the avian sex-determination mechanism [6]. Here we show equivalent expression of at least six of nine Z chromosome genes in male and female chick embryos by using real-time quantitative PCR [7]. Only the Z-linked ScII gene, whose ortholog in Caenorhabditis elegans plays a crucial role in dosage compensation [8], escapes compensation by this assay. Our results imply that the majority of Z-linked genes in the chicken are dosage compensated.

DNA methylation is essential for murine development and is implicated in the control of gene expression. MeCP2, MBD1, MBD2, MBD3, and MBD4 comprise a family of mammalian, nuclear proteins related by the presence in each of an amino acid motif called the methyl-CpG binding domain (MBD). Each of these proteins, with the exception of MBD3, is capable of binding specifically to methylated DNA. MeCP2, MBD1 and MBD2 can also repress transcription. We describe the genomic structure and chromosomal localization of the human and murine Mbd1, Mbd2, Mbd3, and Mbd4 genes. We find that the highly similar MBD2 and MBD3 proteins are encoded by genes that map to different chromosomes in humans and mice but show a similar genomic structure. The Mbd1 and Mbd2 genes, in contrast, map together to murine and human Chromosomes (Chrs)18. The Mbd3 and Mbd4 genes map to murine Chrs 10 and 6, respectively, while the human MBD3 and MBD4 genes map to Chrs 19 and 3, respectively.