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Dissecting the ultrastructure of meiotic chromosomes and the role of the chromosome periphery using advanced imaging
Get PDFUnderstanding the three-dimensional structure of chromosomes is essential for elucidating the mechanisms that govern their behaviour during cell division. However, detailed ultrastructural analysis of chromosomes in oocytes has been limited by technical challenges associated with imaging large, spherical cells. This thesis describes the development and application of a three-dimensional correlative light and electron microscopy (3D-CLEM) pipeline to overcome these challenges and enable high-resolution visualisation of chromosomal architecture during meiosis.
The 3D-CLEM workflow was first established and optimised using mitotic cells, allowing for precise correlation between fluorescence and electron microscopy images. This approach led to the generation of a high-resolution 'mitotic atlas', enabling the structural classification and quantitative comparison of mitotic chromosomes. Notably, this work contributed to the identification and characterisation of a novel chromatin state—plateaud compaction—recently reported in Cisneros-Soberanis & Simpson et al., (2024).
Following successful implementation in mitotic systems, the pipeline was applied to mouse oocytes to investigate the organisation and function of the chromosome periphery during meiosis. Immunolabelling and correlative imaging confirmed the presence of the proliferation marker Ki-67 at the periphery of meiotic chromosomes, establishing its role as a key organiser of this structural domain. Targeted disruption of the periphery revealed its critical importance for maintaining chromosome integrity, with its removal resulting in altered chromatin morphology and disorganisation within the meiotic spindle. This exciting discovery led to a second publication currently in pre-print and under review with Nature Comms (Simpson et al., 2024)
Finally, the pipeline was extended to the study of human oocytes, where it enabled the first three-dimensional reconstructions of meiotic chromosomes. These early datasets demonstrate the challenges associated with analysis of these cell types and defines key characteristics to make this work possible. Following development of the technique for handling human oocytes the feasibility of 3D-CLEM for human oocyte research was proven and the data provides a foundation for future investigations into chromosomal architecture in human meiosis.
In summary, this work establishes and validates a novel 3D-CLEM approach for high-resolution structural analysis of chromosomes in large mammalian oocytes. The pipeline has been successfully applied to both mouse and human cells, yielding new insights into the role of the chromosome periphery and enabling enhanced visualisation of chromosome structure during meiosis. This methodological advancement opens new avenues for investigating the ultrastructure of chromosomes and associated sub-nuclear compartments in rare or challenging cell types
A critical philosophy of microaggressions
Get PDFThis thesis addresses three main questions: (1) ‘what are microaggressions?’; (2) ‘how do they harm marginalised people?’; and (3) ‘how should we deal with them?’.
I begin by offering a brief review of the psychological and philosophical literature on microaggressions, with a particular focus on the critique that the microaggression concept is too ill-defined to offer certainty over what counts as a microaggression. I intervene in this debate by rejecting the claim that we require this level of certainty and defend a structural account of microaggressions which characterises them as a subtly oppressive social practice. Following this, I motivate why microaggressions are oppressive by exploring the ways in which they constitute a form of violence against marginalised people. More specifically, I introduce the concept of existential violence to describe how microaggressions diminish the subjectivity of their targets by denying, homogenising, and inferiorising their identities and experiences, and substantiate this by offering a phenomenological account of microaggressions.
To address how we should deal with microaggressions, I first consider how we might respond to the microaggressor. Starting with anger, I warn against uncritically buying into the argument that anger is a ‘counterproductive’ response to microaggressions by exposing how this critique promotes a masterful form of disciplinary control which reinforces the oppression of marginalised people. Next, I explore the potential for disarming microaggressions through cringe, putting forward my own account of ‘cringeworthiness’ in order to show how microaggressions can be understood as cringeworthy, and outlining some of the benefits and drawbacks of cringing at microaggressors. Finally, I consider what each of us can do to reduce the likelihood that we engage in microaggressions. Moving beyond popular rhetorics around acknowledging our bias and privilege, I argue that we should embrace a politics of disorientation, characterised by an openness to being transformed by those moments of discomfort where our habits of oppression, like microaggressions, become disrupted
Teaching children to be Soviet: a textual and visual analysis of Murzilka’s role models in the era of de-Stalinisation.
No full textThe focus of this thesis is the Soviet children’s magazine Murzilka in the period 1956 – 1964 and the role models presented therein. From its very foundation, children were viewed as vital to the success of the Soviet Union. Literature as a means of guiding, educating and shaping these children became of paramount importance to the regime. Murzilka was one of the earliest magazines for Soviet children, and is now recognised as the longest-running in the world.
By the 1960s, by its own estimation, Murzilka boasted a readership of twenty million children. In the years after the death and denunciation of Stalin, how did the magazine speak to these young readers? What were the main themes of its content? In particular, with the dismantling of Stalin’s personality cult, what role models did the magazine offer its readers in the era of de-Stalinisation? How did Murzilka portray these role models, and what norms and values did these embody and enact for children to emulate? In order to tackle these questions, this study examines closely the visual and textual content of Murzilka at this particular moment in time, relating its analysis to themes and tropes more widely prevalent in propaganda and culture of the Thaw period.
This study looks at the written and illustrated content of the magazine both quantitatively and qualitatively. Initial examination of the magazine in the period 1956 – 1964 identified a number of recurring role models: Lenin, the military, parents and cosmonauts all featured frequently on the pages of the magazine. Closer reading of the source material and deep content analysis seeks to understand how these recurring role models were presented to readers and how they were exploited as a means of modelling appropriate behaviour.
This thesis provides a focused case study of four major role models presented to Soviet children on the pages of Murzilka in the period 1956 – 1964. Through careful, in-depth content analysis, I consider change and continuity in the literary and visual representation of these role models in relation to broader socio-political and cultural developments. This contributes to a better understanding of the ideological, cultural, and moral underpinnings of de-Stalinisation
Hydrogel-based local delivery loaded with Poly(β-Amino Ester) (PBAE)-siRNA polyplexes for glioblastoma therapy
No full textGlioblastoma (GBM) is an aggressive and highly heterogeneous primary brain tumour known for its resistance to standard therapies and frequent recurrence, particularly during the critical gap period between surgical resection and the initiation of chemoradiotherapy. Gene therapies using small interfering RNA (siRNA) have emerged as a potential approach to selectively silence genes involved in cancer progression. However, the clinical application of siRNA remains limited due to its instability, inefficient delivery to target tissues, off-target effects, and the difficulty of achieving delivery within the brain cavity. To overcome these limitations, this study aims to develop and evaluate a hydrogel-based local delivery system incorporating poly(β-amino ester) (PBAE) polyplexes for targeted and sustained siRNA delivery, to control GBM progression during the critical therapeutic gap window. In this study, both linear and branched PBAEs were synthesised via Michael addition reactions using various acrylate and amine monomers. Among the formulations, the branched BF-TMPTA20 PBAE demonstrated the most promising performance. Gel permeation chromatography (GPC) analysis revealed a molar mass of 10200 Da, and NMR confirmed successful end-capping. When complexed with scrambled siRNA, BF-TMPTA20 polyplexes maintained a positive surface charge and particle sizes below 200.0 nm across all polymer:siRNA w/w ratios, with the strongest binding affinity observed at a 64:1 ratio. The polyplexes also exhibited non-toxicity in GIN31-fluc cells and slightly higher transfection efficiency compared to Lipofectamine-3000 at the optimal w/w ratio. Cellular internalisation studies in both 2D and 3D cell culture revealed efficient intracellular delivery and escape from endosomal/lysosomal compartments, indicating BF-TMPTA20 as a potential carrier for siRNA delivery to GBM cells. This polymer was subsequently complexed with therapeutic siRNAs targeting CDC42BPA and CDC42BPB, genes known to be upregulated in invasive GBM phenotypes.
Knockdown of CDC42BPA and CDC42BPB either individually or in combination significantly reduced GBM cell proliferation and invasion, induced apoptosis, and altered the tumour microenvironment. Dual-siRNA treatment produced significantly greater biological effects than single-siRNA treatments, likely due to complementary or overlapping gene silencing effects. To apply as a local delivery system, the BF-TMPTA20 polyplexes were incorporated into a thermoresponsive mPEG-PCL-HMDI-PCL-mPEG (PECE) triblock copolymer hydrogel. The hydrogel exhibited prolonged siRNA release over a two-week period while maintaining transfection efficiency and minimal cytotoxicity. In conclusion, the BF-TMPTA20 polyplex-loaded PECE hydrogel system offers a promising approach for localised siRNA delivery in GBM. It addresses key limitations in gene therapy, including cellular uptake, endosomal escape, and sustained delivery, and may serve as an effective system to prevent tumour recurrence during the critical post-surgical treatment period
Modelling the dryland agricultural systems of Q’a Shubayqa, Jordan
Get PDFThis project investigates a rainfed agricultural system in the drylands of north-eastern Jordan. The study area, Q’a Shubayqa, is characterized by very low rainfall rates but supports a cereal-based mono-cropping system. Local farmers were surveyed to define the main components of the agricultural system and its limiting factors. Farmers confirmed that system productivity has been affected by climate change and severe weather events during the last three decades. Results of the farmer survey and remote sensing and climate data analyses have shown that growth and development of cultivated wheat and barley crops in Q’a Shubayqa mainly depend on seasonal flash-floods that occurs following winter rainfall in the south-western highlands of Syria. Hydrological SWAT modelling of the study area watershed, undertaken for a long-term period to quantify flood water has shown unstable patterns in terms of quantity and timing which would affect cereal planting dates. Historical grain yield simulations using genetic coefficients of the studied cultivars, that were derived in this study from observed data collected from different environments across Jordan and over different seasons, have shown that planting date is a limiting factor for both crop types. Using these DSATT models the impact of different climate change scenarios through to 2100, on grain yield and grain filling period was investigated. Most of studied cultivars were sensitive to future climate change where grain yields decreased, and grain filling period reduced. The effect of high atmospheric CO2 levels on grain yield was limited with a slight increasing but gain yield still below the grain yield baseline. The impacts of additional agricultural practices were evaluated when simulating grain yield under future climate change considering high atmospheric CO2 levels and results showed that delayed planting date has decreased grain yield, however organic amendment positively affected grain production. Results suggest that wheat cultivar Hourani and barley cultivar Rum are less affected by climate change
The application of machine learning to understand the dynamics of soil structure
Get PDFMachine learning and artificial intelligence are quickly changing the world. Just a few years ago, these concepts usually provoked visions of theoretical fiction. However, now they are becoming commonplace across a vast array of industries and even in our daily lives.
Soil structure is a very important factor in soil health and has strong applications in agricultural studies and improvement. It is also a dynamic, complex, and very hard-to-measure component. Scientific advancements which improve the accessibility of soil structure quantifications are of great importance and benefit to the soil science community.
X-ray Computed Tomography (X-ray CT) is a non-destructive and non-invasive technique for observing and analysing soil structure. It has been used in soil science for several years for various tasks, leading to a large volume of data to be analysed. A lot of this data is often discounted, due to the extreme volumes produced.
Machine learning allows for high-throughput data processing, unrestricted by human processing and labour limitations. Data processing and analysis via machine learning algorithms might also notice new trends or patterns in the data that would not be noticed with human analysis. This means that machine learning algorithms could be the perfect solution to take a deeper dive into the data collected from X-ray CT, get more information out of the data, and potentially find new patterns.
Soil structure can vary greatly due to the soil composition, weather conditions (temperature, rain, or lack thereof), crops, compaction, tillage management and the biodiversity in the soil. In recent years there has been a lot of interest in zero tillage, as part of a series of measures usually deployed under the banner of conservation or regenerative agriculture and how it may be beneficial over conventional tillage. This research looks at how the tillage practices impact the soil structure and in particular the pore network, and how these properties may vary over a growing season. Different machine learning techniques are looked at and applied to demonstrate how machine learning can be used alongside X-ray CT images and soil data to improve processes and observe new information.
Ultimately, the large volume of data produced by X-ray CT make excellent training datasets for machine learning algorithms. The neural networks demonstrated were mostly successful at identifying and predicting soil type, water content and tillage treatment based on soil structure. Shortcomings could be overcome by increasing the volume and variation of training data, as well as making use of more advanced machine learning techniques such as multi-resolution networks
Probing USP4 Specificity through Engineered Substrate Trapping and a Fluorescence Polarization Assay Platform Suitable for Measuring DUB Activity and Inhibition
No full textDeubiquitinases (DUBs) play a critical role in the regulation of various cellular processes, such as protein homeostasis and signaling, rendering them attractive drug targets. However, the generation of reagents for measuring DUB activity typically involve several steps and is not straight forward. To dissect their activity and substrate recognition. Here, we report the development and characterization of a novel fluorescent polarization assay using an isopeptide bond substrate mimetic (IsoMim) that can be made recombinantly in high yields. The IsoMim assay was able to discern the differential activity of ubiquitin specific protease family members (USP4, USP15, USP11 and USP2), the ubiquitin C-terminal hydrolase UCHL3 and the Machado-Joseph Domain deubiquitinase JOSD2. A competition assay format of the assay was developed that discerned differences between the close paralogues USP15, USP4 and USP11 in interacting with monoubiquitin, the isopeptide mimetic ubiquitin-GGG and the C-terminal truncation variant ubiquitin (1-74). Moreover, dose-response curves and associated pIC50 values using the broad-spectrum inhibitor PR-619 confirmed differential inhibition in the low µM range for four tested DUBs, demonstrating assay reliability. The successful discrimination of DUB activity and inhibition, and the high obtainable yields of the substrate make the IsoMim assay method applicable for high-throughput screening (HTS). This was ascertained in a ‘pseudo HTS screen’ for USP4 inhibitors in which PR-619 was successfully identified as a ‘pseudo hit’. USP4 was then used to dissect the molecular basis of substrate recognition and specificity, we determined high-resolution crystal structures of USP4 catalytic core (D1D2) bond to both linear diubiquitin and an engineered Ub-GG-Ub mimetic, revealing an induced-fit mechanism in which blocking loops (BL1 and BL2), switching loop (SL) and finger domain undergo concerted rearrangements to accommodate diubiquitin. A potential secondary S2 binding site on the finger domain was identified in the Ub-GG-Ub complex. Mutagenesis of S2-localized USP4 residue His805 to glutamate (H805E) and subsequent cleavage assays showed minimal impact on activity against linear tetraUb and K63-linked polyUb. We also examined a critical gate keeper residue mutation localized in BL2 S1’ site, V879D in blocking loop 2 (BL2), selectively impacted USP4’s ability to cleave linear diubiquitin, as confirmed by both gel-based assays and FP analysis. To further probe the S1’ site and assess substrate preferences, we developed a series of monoubiquitin based FP probes with distinct C-terminal extensions, Ub-GGGC-FM, Ub-MGGC-FM, and Ub-MQIC-FM. These tools enabled quantitative profiling of USP4 specificity and revealed altered binding behaviors upon mutation of conserved S1’ site residues His369 and Asn901.
Altogether, the developed assay provides a valuable tool for probing DUB activity and the identification and characterization of DUB inhibitors and has the potential to accelerate drug discovery efforts in this area. Our study uncovers key structural and functional features that govern USP4’s substrate recognition, defines roles of auxiliary binding sites and loop regions, and introduces versatile FP probes suitable for mechanistic and screening applications
A feasibility study for a randomised control trial on the effects of engagement with a dental anxiety online peer support group
Get PDFThe study presented in this thesis aimed to evaluate the feasibility of a Randomised Control Trial (RCT) on the effects of engagement with a Dental Anxiety (DA) Online Support Group (OSG) in an adult population. The purpose of a feasibility study is to ascertain whether an RCT should be conducted, can be conducted, and if so, how it should be conducted.
Background
DA is a common phenomenon, with an estimated 54% of the population in England reporting moderate to extreme anxiety. It is a condition that is more likely to affect women, younger adults and people from lower socio-economic backgrounds. DA can lead to avoidance of routine dental appointments, shame and stigma, and poorer oral health outcomes, and this cycle can be self-perpetuating. There is evidence that the impact of DA can be wide-ranging, negatively affecting self-esteem, health-related quality of life, eating habits and even relationships. DA often starts in childhood and a variety of exogenic and endogenic factors have been suggested as potential causes. Ways of managing DA currently include pharmacological (for example sedation or analgesia) and psychological therapies (for example hypnosis or cognitive behavioural therapy). However, pharmacological therapies only address the symptoms and access to psychological therapies is limited.
In the context of healthcare, OSGs are virtual communities, often focused on a specific condition or illness, where peers can share their lived experience, exchange information, provide and receive both emotional and practical support. OSGs can use a variety of platforms (for example chat rooms, discussion forums or social media sites) and can be either synchronous or asynchronous, moderated or unmoderated. People who live with conditions that are rare or that carry stigma are more likely to be motivated to join an OSG, and the anonymous nature of OSGs may encourage disclosure. Most OSG users are readers (commonly referred to as lurkers in the literature) rather than active posters. Some of the potential negatives of OSGs are misinformation, negative content and negative online behaviours. There is some evidence that OSGs can support behaviour change and therefore promote health outcomes, however much of that evidence is qualitative, based on a self-selecting population of existing OSG users, or has methodological issues.
Systematic Review
Systematic reviews can help assess whether an RCT should be carried out on a specific topic. Therefore, a qualitative systematic review was conducted in accordance with JBI methodology to explore the experiences of users of OSGs for adults living with DA. Rigorous searches were conducted across seven databases but yielded only 2 papers. This limited number of papers shows that OSGs for DA are an under-researched area. Both studies were considered of sufficient quality to be included in the review but had some methodological limitations. Meta-aggregation led to five synthesised findings, most of which were rated as moderately dependable. These synthesised findings indicate that participation in an OSG can be a positive experience for adults with DA and help bring about positive cognitive and behavioural changes.
Methodology
Pragmatism is the chosen paradigm for this feasibility study, along with a concurrent mixed-methods design (where qualitative and quantitative data are collected in parallel) as this reflects the complexity of the topic and allows participants’ perspectives to be considered. Both pragmatism and mixed methods are well-suited to a feasibility study.
The protocol was developed with input from a Patient and Public Involvement (PPI) focus group and panel, who provided input on their lived experience of DA and who gave feedback on a draft protocol as well as on the participant-facing materials (e.g. advert, questionnaire, participant information sheet). Their input was taken forward and shaped the final protocol that was granted approval by the University of Nottingham’s Faculty of Medicine and Health Sciences’ Research Ethics Committee in April 2023.
The study aimed to recruit 38 adults with moderate to high DA, as measured by the Modified Dental Anxiety Scale (MDAS), within 3 months. This sample size was based on a review of the literature on sample sizes for feasibility studies and allowed for an attrition rate up to 33%, based on evidence that online interventions can have high attrition. Recruitment took place through social media, predominantly through Facebook. Participants were randomly allocated to either a Wait-List Control (WLC) condition or to the OSG condition and asked to use it in a naturalistic manner (with no frequency or amount of time mandated) for a period of six weeks. Their usage of the OSG was measured by capturing logon data daily. At the end of the six weeks, semi-structured interviews were conducted with participants allocated to the OSG to explore the acceptability of the OSG intervention. Thematic analysis was conducted on the transcripts from the interviews. Participants completed pre- and post- measures of DA (MDAS) and intention to attend routine dental care (Dentist Contemplation Ladder). Descriptive statistics were produced to analyse the data.
Results:
40 participants were recruited within 3 months and attrition was lower than anticipated at 7.5%. Reductions in the mean MDAS score (indicative of a reduction in DA) were seen in both the OSG and the WLC conditions but were greater in the OSG condition. Increases in the mean Dentist Contemplation Ladder (indicative of a greater readiness to attend an appointment) were seen in both the WLC and the OSG conditions but again were greater in the OSG condition. The effect size based on the MDAS indicates that a sample size of 72 would be required for a fully powered RCT.
Thematic analysis generated 4 themes: “Initiation” which examines the barriers and facilitators to accessing the OSG, “exploration” or how participants navigated the OSG, “engagement” and what fostered or hindered participants participation in the OSG, and finally “benefits and outcomes” that the OSG had fostered. Most participants were positive about the experience, with a small number rejecting the OSG, having decided early on that it did not meet their needs.
From a process perspective, there was variation in the level of usage of the OSG. Overall, the number of times participants logged on to the OSG was limited with a mean of 3.6 times over six weeks. Participants found the sign-up process and the measures simple and easy to complete. The phone or Teams call that formed part of the sign-up process was well received by participants and may have helped reduce the risk of imposter participants.
Conclusions:
The results of the study indicate that it is feasible to recruit sufficient eligible participants for an RCT after taking attrition into account. The usage of the OSG was acceptable to the participants in this study, although some participants engaged more than others. The MDAS would be an appropriate main outcome measure for an OSG, although both measures provided useful insight and were acceptable to participants. A full RCT is therefore both feasible and necessary to further understand the effect of OSGs on DA
“Black, Puerto Rican, lesbian literature”?: representations of Yolanda Arroyo Pizarro
Get PDFThis thesis analyses representations of author Yolanda Arroyo Pizarro (Guaynabo, Puerto Rico, 1970) and her literary work as Black, Puerto Rican, and lesbian. It questions why and how the identity categories of race, sexuality, gender, and nationality are foregrounded in representations of this author and her body of work, while considering the implications of this categorisation process for literature more broadly.
Where previous scholarship on Arroyo Pizarro has centred textual analysis, this thesis offers a new approach by focusing on extratextual representations. The thesis takes much inspiration from the relatively small body of reader reception studies of live reader participants, as well as research on paratexts.
Stuart Hall’s work on representational systems provides a framework for this study of literary categories. Analysis incorporates paratextual and metatextual sources such as reviews, criticism, and social media posts, which give an initial picture of the ways in which Arroyo Pizarro and her texts are represented. Following this, the focus moves to analysis of interviews conducted with fourteen participants including the author, book industry professionals, academics, and other readers. This interview methodology produced new representations of Arroyo Pizarro and her work from a variety of perspectives. While the sample is small and cannot be considered representative, it offers rich, qualitative data on participants’ personal relationships with this author, her writing, and the phenomenon of identity-based literary categories.
Analysis of interview data centres on participants’ representations of the author, her texts, and her readers. Participants foregrounded race, sexuality, gender, and nationality in their representations of Arroyo Pizarro and her work, representing both as Black, Puerto Rican, and lesbian. Terms like “Black literature” and “literatura lésbica” are ultimately found to resist strict definitions. Prompted to take positions on the political implications of categorising literature by identities, participants acknowledged the potential for such categories to be reductive but generally argued that their use is necessary for the visibility and empowerment of marginalised groups. On the question of readers, interview participants constructed age, politics, and religion as more important factors than race, gender, or sexuality in determining whether an individual would read and enjoy Arroyo Pizarro’s work.
The thesis contributes an innovative approach and methodology within the field of literary studies. The focus on extratextual representations demonstrates that literary texts are not autonomous, discrete objects, but are produced by discourses of power and identity
Engineering biomimetic in vitro models for personalized cancer treatments
No full textColorectal cancer (CRC) is the second leading cause of cancer-related deaths worldwide. Conventional therapies often fail due to the inability of current in vitro models to recapitulate the complexity and heterogeneity of the tumour microenvironment (TME). Most research still relies on 2D cultures, which poorly model tumour biology or on 3D models that use animal-derived matrices such as Matrigel, which suffer from high variability and limited tuneability. To address these limitations, this thesis presents the development and optimisation of peptide amphiphiles (PA)-based composite hydrogels co-assembled with tumourspecific CRC-ECM macromolecules (PA-ECM), including collagen, hyaluronan, fibronectin, and laminin. By engineering these nanofibrillar hydrogels with either random (rPA) or aligned (aPA) architectures, it is possible to closely mimic the CRCTME, enabling precise control over ECM composition, nanofibrillar organisation, and mechanical properties. Structural, biochemical, and mechanical analysis demonstrated that rPA-based hydrogels enable a more stable and homogenous ECM incorporation, maintaining a physiological soft stiffness. In contrast, aPA-based hydrogels were stiffer, less permissive to ECM incorporation, and failed to support long-term cell viability. Using CRC cell lines and patient-derived organoids (PDOs), we demonstrate that rPA-based hydrogels support long-term cell viability and growth, preserving tumour-specific morphology and gene expression profiles of primary colorectal tumours. Transcriptomic analysis revealed that ECM incorporation in rPA hydrogels modulates gene expression related to ECM remodelling, cell adhesion, and signalling pathways key in CRC progression, while maintaining lineage-specific differentiation crucial for disease modelling. In order to achieve a physiologically relevant model, rPA-based hydrogels were tested in immune-tumour co-culture models. These hydrogels maintained structural integrity, promoted CD8⁺ T-cell infiltration, and enabled the assessment of chemotherapeutic and immunotherapeutic responses. By integrating tumour-specific ECM macromolecules with architecturally defined PA nanofibres, this platform bridges the gap between simplistic 2D in vitro models and clinical 3D tumour complexity. This approach offers a complex, yet defined and reproducible platform with significant potential to not only recapitulate CRC-TME but also a wide range of different types of other TMEs
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