research paper on plant based meat

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Plant-based and cell-based approaches to meat production

Natalie r rubio, david l kaplan.

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Corresponding author.

Received 2020 Aug 4; Accepted 2020 Nov 6; Collection date 2020.

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/ .

Advances in farming technology and intensification of animal agriculture increase the cost-efficiency and production volume of meat. Thus, in developed nations, meat is relatively inexpensive and accessible. While beneficial for consumer satisfaction, intensive meat production inflicts negative externalities on public health, the environment and animal welfare. In response, groups within academia and industry are working to improve the sensory characteristics of plant-based meat and pursuing nascent approaches through cellular agriculture methodology (i.e., cell-based meat). Here we detail the benefits and challenges of plant-based and cell-based meat alternatives with regard to production efficiency, product characteristics and impact categories.

Subject terms: Synthetic biology, Tissue engineering, Environmental impact, Sustainability

Large-scale meat production can have negative impacts on public health, the environment and animal welfare. In this Review, the authors consider plant-based and cell-based approaches to meat production and the challenges they face.

Introduction

Global production and consumption of meat continue to surge as demand is driven upward by population growth, individual economic gain, and urbanization 1 , 2 . In 2012, the Food and Agriculture Organization (FAO) of the United Nations projected the global demand for meat would reach 455 M metric tons by 2050 (a 76% increase from 2005) 3 . Likewise, the global demand for fish is projected to reach 140 M metric tons by 2050 4 . The majority of this incline is attributed to middle-income countries (e.g., China), as consumption in higher-income countries is relatively stagnant or marginally decreasing (e.g., United Kingdom) and in lower-income countries, the rate of consumption is fairly constant (e.g., India) 1 . This pattern is consistent with a proposed theory that the relationship between meat consumption and income follows an “inverted U-shaped” trend; consumption initially increases with rises in income but eventually reaches a turning point at which consumption stagnates or declines 5 . This observation may be rationalized by correlations between high income and increased concern for the consequences of animal agriculture 5 .

This rising demand is problematic as current methods of large-scale animal husbandry are linked to public health complications, environmental degradation and animal welfare concerns. With regard to human health, the animal agriculture industry is interconnected with foodborne illness, diet-related disease, antibiotic resistance, and infectious disease 6 , 7 . Notably, zoonotic diseases (e.g., Nipah virus, influenza A) are linked to agricultural intensification and meatpacking plants in the United States were hotspots for COVID-19 outbreaks 7 , 8 . Animal agriculture also contributes to environmental issues including greenhouse gas emissions, land use, and water use 1 . The United Nations Intergovernmental Panel on Climate Change released a 2018 report asserting that greenhouse gas emissions must be reduced 45% by 2030 to prevent global temperatures from increasing 1.5 °C; a target that could mitigate catastrophes associated with a 2.0 °C increase 9 . Conventional mitigation techniques include improvements in reforestation, soil conservation, waste management as well as tax policy, subsidies, and zoning regulations 10 . While these strategies remain important, the urgency of climate change may require more transformative approaches. Lastly, with regard to animal welfare concerns, each year billions of animals are killed or suffer either directly (e.g., farm animal slaughter, seafood fishing) or indirectly (e.g., fishing by-catch, wildlife decline due to habitat destruction) in relation to human food systems 11 , 12 .

The majority of the aforementioned issues can be attributed to the fact that the raw material inputs (i.e., animals) for conventional meat production are inherently unsanitary, inefficient, and sentient. By removing animals from the manufacturing process, several externalities may be alleviated. Plant-based meat (PBM) and cell-based meat (CBM) approaches offer to generate food from non-animal sources. While traditional PBMs (e.g., tofu) have existed for centuries, novel PBM alternatives with enhanced sensory characteristics have been commercialized more recently (Fig.  1 ) 13 . Other groups have initiated the development of a new field—cellular agriculture. Cellular agriculture describes the tactic of producing commodities from cells, rather than whole organisms or animals, such as CBM; meat grown from muscle or fat cells rather than cows, pigs or chickens. While there is evidence to support certain benefits of these approaches compared to animal-based meat (ABM), it is important to more comprehensively evaluate impacts on human health and the environment as production systems evolve. In addition, widespread adoption of these products will require more direct benefits to the consumer; such as taste, cost, and convenience 14 . This review serves to compare plant-based (i.e., meat analogs composed of plant proteins) and cell-based (i.e., meat analogs generated from cell cultures) methods to educate stakeholders about the strengths and challenges of each approach and highlight areas of uncertainty.

Fig. 1. The history and evolution of animal-, plant- and cell-based approaches to meat production.

13 , 87 – 93 . Humans have consumed plant-based meat (2555 years ago) for only 0.098% of the time period for which their ancestors have consumed animal-based meat (2,600,000 years ago). Likewise, humans have eaten cell-based meat (7 years ago) for only 0.274% of the time period for which they have consumed plant-based meat.

History and approach

Plant- and fungi-based meat (PBM) products encompass the flavor, texture, and/or nutritional aspects of meat but are different in composition; namely are made from non-animal sourced materials. Based on the time of development and technical complexity, PBM products can be differentiated into two flexible categories: traditional and novel (i.e., next-generation) 13 . Traditional meat analogs were developed thousands of years ago in Asia and include relatively simple derivatives from soybeans (i.e., tofu, tempeh) or wheat (i.e., seitan) 13 . In contrast, novel PBMs are characterized by the design and marketing of products as near equivalent replacements for ABM with regard to taste, texture, and nutrition. Product categories can also exist between traditional and novel, as they may meet some but not all of the aforementioned criteria. A distribution map of global companies and brands developing novel PBMs can be found in Fig.  2 . Typically, the production of PBM includes three steps 15 : (i) Protein isolation and functionalization—Target plant proteins are extracted from plants, some of which are subjected to hydrolysis in order to improve their functionalities such as solubility and cross-linking capacity; (ii) Formulation—The plant proteins are mixed with ingredients to develop meat texture such as food adhesives, plant-based fat and flour. Nutrients are added to match or exceed the nutrient profile of the meat. (iii) Processing—The mixture of plant proteins and other ingredients undergo protein reshaping processes (e.g., stretching, kneading, trimming, pressing, folding, extrusion, etc.) to form a meat-like texture. Innovative technologies being utilized to improve the organoleptic properties of PBMs include shear cell technology, mycelium cultivation, 3D printing, and recombinant protein additives 16 , 17 .

Fig. 2. Geographical distribution of plant-based (green circles) and cell-based (orange circles) meat companies.

Companies were included as listed in the Good Food Institute alternative protein company database (August 2020).

CBM, also referred to as in vitro meat, lab-grown meat or cultured meat, is meat produced by cultivating cells as opposed to farming animals. CBM technology is based on advances in stem cell biology (e.g., induced pluripotent stem cells) and tissue engineering (e.g., in vitro skeletal muscle grafts) originally purposed for medical applications. CBM production involves four central components: (1) muscle and fat cell isolation and culture, (2) xeno-free culture medium formulation, (3) scaffold development, and (4) bioreactor design; the details of which are described extensively elsewhere 14 . Interestingly, the concept of CBM can be traced back to 1930 when Frederick Smith, the British Secretary for India, envisioned the genesis of “self-reproducing steaks” through an excerpt of his essay collection The World in 2030 AD , which reads: “It will no longer be necessary to go to the extravagant length of rearing a bullock in order to eat its steak. From one ‘parent’ steam of choice tenderness, it will be possible to grow as large and as juicy a steak as can be desired 18 .” While CBM has yet to be commercialized in 2020, noteworthy progress has taken place over the past couple of decades. Key milestones include the first CBM patent filed by Willem van Eelen in 1999 19 , the first peer-reviewed research on cultured fish funded by NASA in 2002 20 and the first cultured beef burger debuted by Maastricht University in 2013. Today, there are dozens of start-up companies around the globe working to bring CBM products to market.

Key sources of protein inputs for novel PBM are relatively inexpensive. The majority of plant-based products are primarily formulated with pea, soy or wheat protein. The agricultural prices (received by farmers in the United States) for these key proteins are 3.8–12.7 times lower than prices received for cattle, hogs, and broilers. In fact, sometimes soy and wheat are combined with ABM to reduce costs for processed meat products 21 . When standardized by cost (2009 data) per gram of protein, soybeans ($0.01/g) and wheat ($0.03) are still remarkably less costly than cows ($0.32/g), pigs ($0.22/g), and chickens ($0.12/g) 22 . Despite this glaring discrepancy on the key protein input level, novel PBMs tend to cost more than their animal-based counterparts in a retail setting. This discrepancy may be partially due to processing costs as 94.3% of retail costs for crop products are associated with post-harvest processes, while this accounts for ~50% of the retail costs for beef 22 . Furthermore, aside from primary proteins, PBMs often include plant-based fats, flavor enhancers, and color additives which contribute to cost. Some consumers are willing to pay premiums for meat substitutes linked to personal health benefits 23 .

Economic feasibility is a significant hindrance to CBM commercialization 24 . The cultured beef burger cultivated by Maastricht University in 2013 is reported to have cost $280,400 ($2,470,000/kg) to produce. The production process involved three researchers using bench-scale techniques to culture 20,000 muscle fibers over three months and served as a proof-of-concept rather than an attempt to scale production. A few groups have performed preliminary economic analyses to project the cost of CBM for large-scale production scenarios (Table  1 ). In 2008, The In Vitro Meat Consortium estimated, by modeling capital and growth medium costs based on data for single-cell protein production, CBM could cost approximately twice as much as chicken 25 . In 2014, a study speculating on the technical, societal, and economic factors of village-scale CBM production calculated a cost range of $11–520/kg dependent on the price of growth medium 24 . Invertebrate (e.g., insect, crustacean) cell culture may present a more cost-efficient platform for CBM production based on the unique properties of insect cells (e.g., xeno-free growth medium, high-density suspension culture) 26 , 27 . Select companies are targeting high-value products (e.g., foie gras, bluefin tuna, kangaroo meat) in order to lower the bar for reaching price parity. Interesting, a recent consumer acceptance study from the Netherlands reported 58% of participants were willing to pay a 37% premium for cell-based beef compared to conventional animal-based beef 28 .

Results from preliminary economic analyses of CBM production 24 , 25 .

a Estimates are based on capital, variable media, and overhead costs for a plant capacity of 15,000 tons/year. The lower limit is estimated for cells grown in suspension and the upper limit is estimated for cells grown in a 3D matrix.

b Estimates are based on batch production of 2560 kg CBM per batch assuming 20,000 L/batch. CBM cost only considers contributions from growth medium.

Regulatory framework

PBMs are regulated in a similar manner as other non-animal foods. In the United States, the Food and Drug Administration (FDA), and specifically the Center for Food Safety and Applied Nutrition (CFSAN), oversees food inspection, labeling, packaging, imports, and facility safety. Most PBM products contain simple ingredients that have previously been approved for human consumption. Novel ingredients may be subject to additional evaluation processes. For example, soy leghemoglobin, produced via genetic engineering, filed for “generally recognized as safe” status with the FDA for use as a color additive 17 . In the European Union (EU), current policy and regulation are supportive for alternative proteins innovation and investment. In 2018, the European Commission presented a “EU Protein Plan”, which encourages the production of alternative proteins for human consumption, and listed existing EU policy instruments that “provide options for strengthening the development of EU-grown plant proteins”. Many novel PBM products are classified under the Novel Food Regulation which regulates “food that had not been consumed or do not exist in the EU before 15 May 1997” 23 . Australia, Canada, and New Zealand have also introduced legislation to guide oversight of novel foods 13 . Government oversight is also required for food labeling. In 2018, The United States Cattlemen’s Association petitioned the Food Safety and Inspection Service (FSIS) “to exclude products not derived directly from animals raised and slaughtered from the definition of “beef” and “meat” 29 . The use of terms such as steak, sausage, bacon, fillet, etc. for PBMs is subject to scrutiny and restriction in many EU member states as well.

Oversight for CBM involves the regulation and monitoring of production, packaging, labeling, and marketing. In the United States, CBM will be jointly regulated by the FDA and the United States Department of Agriculture (USDA) based on a decision announced by the departments in 2019 30 . The FDA will regulate cell isolation, storage, growth, and maturation. After cells and tissues have been harvested, the USDA will monitor products through the remainder of the commercialization process and oversee labeling 30 . Scaffold materials may fall under FDA food additive provisions 31 . Even with a joint effort, it will be important to utilize current systems but also to implement new regulation procedures as the technology continues to advance 32 , 33 . Further complication could arise if companies intend to sell products containing genetically modified (GM) cells. While the USDA regulates GM crops, a FDA New Animal Drugs Application provision views DNA manipulation to fall under the definition of a drug and dictates FDA oversight of GM animals; this could potentially be interpreted to also apply to GM cells 33 . A second concern about regulations is with respect to accurate labeling. Similar to the PBM labeling debate, there is an effort to prevent cell-based products from being labeled as “meat” 29 . Based on the Federal Meat Inspection Act which refers to meat as “any product… made wholly or in part from any meat or portion of the carcass”, there may be justification for CBM to retain its wording. In fact, the North American Meat Institute states that cell-based products likely fall into the definitions of either “meat” or “meat byproduct” 34 . For Europe, CBM could be applicable to the European Union Novel Food Regulation pathway. While the Food Safety Authority has approved GM food production, contingent on thorough safety assessments, many European countries (e.g., France, Germany, Greece) have banned the production and sale of GM foods 35 .

Organoleptic properties

The chief organoleptic (i.e., sensory) properties of meat are appearance, aroma, flavor (Table  2 ), and texture. Novel PBMs mimic the look of ABM by manipulating color, fat marbling and structure (Fig.  3 ). Depending on the product, PBMs aim to emulate the appearance of raw (e.g., ground meat) or pre-cooked meat (e.g., deli slices). Heat-stable fruit and vegetable extracts (e.g., apple extract, beet juice) or recombinant heme proteins (e.g., LegH) are used to both recreate the color of fresh meat and change to brown upon cooking 17 , 36 . To mimic the appearance of fat, some novel PBM products exhibit visible semi-solid plant-based fats (e.g., coconut oil, cocoa butter). Engineering flavor and aroma profiles are important to recapitulate the taste and smell of meat. In meat analogs, flavor additives are incorporated to add, enhance or mask specific flavor notes and generally compose 3–10% of the product 21 . Many plant proteins are associated with bitter and astringent tastes, which require selective compound removal by post-processing 15 . In particular, soy products have strong grassy, beany and bitter flavors linked to lipoxygenase, saponin and isoflavone compounds which can be reduced through germination or heating 15 . A synthetic meat flavor developed in the 1980s was composed of sugar, amino acids, nucleotides, glycoprotein, monosodium glutamate, salt, and fat and determined by a sensory panel to be equal or superior to meat extract 37 . Recombinant protein additives like LegH can contribute to the flavor as well as the color of PBMs 17 . PBM texture can be influenced by high-moisture extrusion, shear cell technology, mycelium cultivation and 3D printing. Extrusion, shear cell technology and 3D printing rely on applying mechanical, thermal and shear stresses to a protein mixture to obtain a semi-solid fibrous structure 16 . While many strategies are available to engineer and tune the texture of plant proteins, it can be difficult to balance processing methods to achieve desired mechanical properties while also retaining nutritional value 15 . Conversely, mycelium cultivation involves growing filamentous fungi; particular strains of which emulate the microstructure of meat 38 . Quorn™ is a fungal-based PBM that has provided alternatives for chicken nuggets, meatballs, and minced meat since the 1960s 38 . New start-ups (e.g., AtLast Food Co., Emergy Foods) are growing mycelia with goals of generating higher quality cuts of meat, such as steak.

Precursors and compounds attributable to the aroma and flavor of meat.

Fig. 3. Plant-based and cell-based strategies for emulating appearance properties (color, marbling, structure) of meat.

Structure and marbling are significant contributors to the texture of meat as well as appearance.

To increase the likelihood of mainstream consumer adoption, CBM must be equivalent or superior to ABM from a sensory perspective 39 . The 2013 cultured beef burger (which contained cultured skeletal muscle tissue but not adipose tissue and was flavored with beet juice, bread crumbs, caramel, egg powder, salt and saffron) was described as tasting “like a real burger” by one panelist and “close to meat, but not that juicy” by another 40 . Since this milestone, more effort has been focused on generating cell-based adipose (i.e., fat) tissue; given its significant contribution to taste and texture. Advances in engineering fat tissue for use in food have been reviewed in depth elsewhere 41 . Aside from skeletal muscle and adipose tissue, ABM also contains connective tissue, vasculature networks, and supporting cell types (e.g., fibroblasts). The discrepancies in complexity may result in nuanced flavor differences between ABM and CBM. CBM experts indicate that key flavor profiles can be achieved by co-cultures, medium supplementation, and/or genetic modification 14 , 39 . For instance, researchers have explored the effects of supplementing CBM with extracellular heme proteins (e.g., myoglobin) 42 . Myoglobin is associated with the “bloody” flavor of meat and supplementation was observed to improve the color of CBM constructs without impeding muscle cell growth rates 42 . Early cell-based prototypes emulate processed meat (e.g., burgers, sausages, nuggets) as it is more difficult to emulate the appearance and texture of whole cuts of meat (e.g., steak). As researchers in the field begin to focus on textural properties, significant effort will be required to evaluate myriad factors (e.g., cell to scaffold ratio, the impact of cooking, packaging, storage, and shipping) on tissue structure. CBM texture can be influenced both by cultured cells and supportive scaffolding materials. In vitro skeletal muscle tissue can be engineered to emulate the structure of meat by employing differentiation and cell alignment strategies. For example, mechanical tension, electrical stimulation, and/or micropatterned substrates can be employed to induce cell alignment in vitro 43 . A recent study focused on CBM composed of bovine cells coupled with a textured soy protein scaffold; finding some of the samples exhibited texture (i.e., ultimate tensile strength) properties similar to those of native bovine muscle 44 . In addition, a sensory panel tasted the CBM samples and described “a pleasant meaty flavor” and “a typical meat bite and texture” 44 .

The key plant-based proteins utilized in PBM formulations (e.g., pea, soy, wheat) provide total protein content at levels on par with ABM. However, in order to ensure a balanced amino acid profile, complementation of multiple plant-based proteins is generally necessary. For instance, legume (low in sulfur-containing amino acids, high in lysine) and cereal (low in lysine, high in sulfur-containing amino acids) proteins are favorable complements. Factors that have been identified in plant proteins that may decrease nutrient bioavailability post-ingestion include: structures resistant to proteolysis, protein conformation, and antinutrients (e.g., tannins, phytates, lectins) 45 . Certain processing techniques (e.g., soaking, heating, sprouting) have been shown to increase digestibility 15 . Nutrition is also variable between traditional and novel PBM products. For example, tofu (traditional PBM) and Impossible™ (novel PBM) share certain benefits over ABMs such as containing dietary fiber and minerals while lacking cholesterol. However, tofu-specific benefits include fewer calories, less fat and sodium-free and Impossible™-specific benefits include higher protein and vitamin B 12 content (Fig.  4 ). Concern has been expressed regarding the inclusion of LegH in PBM, citing correlations between heme iron intake and increased risk of diabetes 46 .

Fig. 4. Nutritional value of ABM (beef, pork, and chicken), traditional PBM (tofu), novel PBM (Impossible™ Beef), and mycoprotein (Quorn™) per 100 g wet weight, raw.

Nutritional data for ABM and tofu were obtained from the FoodData Central database (FDC ID: 174036, 167902, 171116, 388713) and Impossible™ and Quorn™ data were obtained from company websites. Content is quantified by the percent of recommended daily intake as determined by the FDA 94 .

Comprehensive, baseline nutrition data for CBM is not publicly available. Using small sample sizes, the nutrient content of cell cultures can be quantified via laboratory assays 27 . Different cell types contribute different sets of nutrients; differentiated muscle cells will likely be the primary source of protein and mature adipocytes can contribute to the fatty acid profile 41 . Certain compounds that are provided by ABM are not present in cultured cells. For example, vitamin B 12 is only synthesized by bacteria and will need to be supplemented 47 . As with flavor, proponents of CBM often claim its nutrition profile will be comparable with or superior to ABM and that nutrients can be tuned via co-cultures, media supplementation, and genetic modification 14 . Media formulation will have a large impact on the viability and efficiency of the cultured cells, on the nutrition profile, and perhaps also impact flavor and taste. Genetic modification for nutritional improvement is another approach that may be more efficient in the long-term, although genetic approaches may pose problems for regulatory strategy and consumer acceptance. Genetic engineering has already been implemented in livestock to improve various aspects of meat production. In 2004, transgenic swine were generated to express a gene originating in spinach with a goal of improving the fatty acid profile of pork 48 . This and other modifications could be implemented on a cellular level to influence the properties of CBM. Comprehensive nutrition data for CBM should become available with the launch of initial products, scale-up, and additional interest from the scientific community.

Consumer acceptance

Consumer acceptance is of particular interest to PBM stakeholders who are looking to increase market share. A high consumer acceptance for PBM products was recorded in China (95.6%) and India (94.5%), compared to the United States (74.7%) 49 . In a European study, the main barriers for dietary inclusion of PBMs were lack of familiarity and low “sensory attractiveness”, and consumers who were unfamiliar with analog products were more likely to want these products to closely imitate ABM 50 . In a focus group study, motivating factors for not eating ABM ranked differently in Germany (e.g., animal welfare, health, environmental impacts), the Netherlands (e.g., animal welfare, poor meat quality, health), and France (e.g., health, animal welfare, sustainability) 51 . For all three nations, the taste was the key factor inhibiting consumption of plant proteins, with other factors including habit, convenience and price. In a sensory panel study comparing animal-, plant- and insect-based burgers, animal burgers were associated with the emotional terms of ‘contented, happy and pleasant’, while plant burgers were associated with ‘disappointed, distrust and discontented’ 52 . Beyond and Impossible™ products introduce a new class of PBM products that more closely mimic ABM compared to the previously established texturized vegetable protein items. These products may be viewed as “highly processed” compared to traditional vegetable burgers and may alienate “clean label” consumers; who are concerned about “unnatural” methods of food production 53 . New consumer acceptance research is required to determine how these products measure up to the findings reported for more established products.

A general consensus in the field is that CBM is targeted at consumers who currently eat meat; as plant-based diets are agreed upon as healthful and sustainable for vegetarian-leaning individuals. Interestingly, vegetarians and vegans in the United States are both more likely to agree with the potential benefits of CBM but less willing to try it compared to omnivores 54 . A systematic review of cultured meat consumer acceptance studies found the most commonly reported concerns were associated with: unnaturalness, safety, healthiness, taste, texture, and price 55 . A 2017 European consumer study found that lack of naturalness decreased acceptance of cultured meat; even with awareness of potential environmental and animal welfare benefits 56 . Along with this finding, research examining internet comments on United States-based news articles covering cultured meat development found more critical input than approval responses, and a frequent critique was that CBM would be “unnatural” and “unappealing” 57 . A 2018 Switzerland study concluded that informing consumers about the production process did not increase acceptance and that communications that emphasize the final product, rather than the technical processes, would be a more successful strategy 58 . Similarly, a 2020 Netherlands study reported that educating consumers on the personal and societal benefits had a positive impact on consumer acceptance 28 . Nomenclature of meat grown from cell cultures may also have an effect on consumer perspective. When comparing the effects of ‘animal-free, clean, cultured’ and ‘lab-grown’ on a panel of participants, ‘animal-free’ and ‘clean’ incited more positive attitudes compared to ‘lab-grown’ 59 . Other common descriptors include ‘artificial’, ‘cell-based’, ‘cultivated’, ‘in vitro’, and ‘synthetic’.

Public health

Meat is an important source of nutrition, especially in developing countries facing nutrition deficiencies. However, the overconsumption of meat has been linked to a number of health concerns. More than 1.8 million people die each year from ischemic heart disease, a quarter of which is linked with overconsumption of certain meat products 60 . Results from a recent clinical trial administered by the Stanford School of Medicine demonstrated that participants who substituted PBM for ABM over eight weeks exhibited lower risk for cardiovascular disease (e.g., reduced fasting serum trimethylamine-N-oxide levels) 61 . The consumption of PBM follows most nutritional dietary guidelines which recommend to limit intake of red and processed meat 62 , 63 , and could benefit consumers that desire reductions in blood pressure, body mass index, and cholesterol 64 . Foodborne pathogens found in meat, such as Escherichia coli, Salmonella , and Campylobacter , result in millions of illnesses each year 65 . Though PBMs are generally not associated with pathogenic disease concerns, non-animal products are capable of causing foodborne illness. A 1999 study screening tofu sold at grocery stores determined that 16% of tested samples were contaminated with coliform bacteria 66 . Plant foods can become contaminated with pathogens via contact with contaminated sources of animal manure, water or other foods. Antibiotics are also used in plant agriculture, but at relatively low levels (in the United States, plant use accounts for only 0.12% of animal agriculture antibiotic use) 67 , therefore, compared with ABM, PBM is less associated with antibiotic-related “drug-resistance” issue.

The commercialization of CBM could impact numerous aspects of public health including foodborne illness, nutrition deficiency, diet-related disease (e.g., colorectal cancer, cardiovascular disease), and infectious disease 6 . The risk of foodborne illness from CBM could be theoretically non-existent, since the sterile conditions required for cell proliferation will prevent contamination with disease-causing pathogens, provided that post-processing and packaging procedures are equally sterile. While sterile cell culture is implemented in pharmaceutical manufacturing, it may not be economically feasible for food production. Natural, food-grade antimicrobial agents may be a promising strategy to reduce contamination risk while remaining cost-effective 68 . Similarly, the threat of zoonotic disease transmission could be directly reduced by decreasing human contact with infected animals and indirectly by reducing habitat destruction 69 . Nutrition deficiency and diet-related disease could be addressed with cell selection, genetic modification, and medium formulation strategies to regulate the presence of healthful and unhealthful compounds. Conversely, consumers report concerns with the safety and healthfulness of CBM, citing fears about unnaturalness, cancer, and inadequate regulation 55 . So far, public health claims on both sides are entirely speculative as the relevant research has yet to be published.

Environmental sustainability

Beyond Meat ® and Impossible™ Foods have both released life cycle assessments (LCA) for their plant-based beef products 70 , 71 . Eutrophication potential and land use requirements for these products are projected to be significantly lower than metrics reported for factory-farmed animal-based beef, pork and chicken while greenhouse gas emissions fall between metrics for pork and chicken and energy consumption exceeds that of pork and chicken (Fig.  5 ). Compared to estimates for Beyond and Impossible™, mycoprotein (i.e., Quorn™) is more impactful with regard to energy and emissions but requires less land for production (Fig.  5 ). The water footprint of PBMs is highly dependent on the source of the main protein. A LCA comparing meat alternatives calculated that mycoprotein-based products (40 kg/kg) have higher water requirements compared to gluten (0.954 kg/kg) and soy-based (0.73 kg/kg) items 72 . A separate LCA study estimated the water usage for 39 distinct meat analogs and determined that, on average, a ton of PBM product consumed 3800 m 3 of water 73 . A majority of the consumption is due to the processing of meat analogs after harvest of the raw protein sources (transportation and packaging were other factors).

Fig. 5. Comparison of the environmental impact of meat and meat analogs.

Data are normalized to the impact of beef production. Eutrophication does not include data for mycoprotein. Land, emissions and energy data for mycoprotein were adapted from a 2015 LCA 72 . Data for beef, pork, chicken and CBM were adapted from a 2015 life cycle assessment 75 . Data for PBM were adapted from an Impossible™ Beef LCA (land, eutrophication, emissions) and a Beyond Meat ® life cycle assessment (energy use) 70 , 71 .

The production of CBM is anticipated to, once optimized, require fewer resources and emit less waste relative to ABM 14 . Anticipated reductions are based on assumptions of: (1) targeted tissue cultivation (i.e., reduced by-products, non-meat tissues); (2) higher production rates and (3) vertical production systems 14 . The first relevant LCA published in 2011 estimated CBM would involve lower energy consumption (7–45%), greenhouse gas emissions (78–96%), land use (99%), and water use (82–96%) compared to ABM 74 . A separate 2015 LCA of CBM reported less dramatic footprint reductions and determined that the energy consumption, acidification potential, and ozone depletion potential impacts of CBM could be more detrimental than ABM, especially when compared to poultry production 75 . CBM is estimated to have a 47% energy feed conversion efficiency and 72% protein feed conversion efficiency, values that are lower than PBM and insect-based meat but higher than ABM 76 .

Animal welfare

PBM products are generally free of animal byproducts and thus do not have direct negative impacts on animal welfare. However, a subset of products contains dairy-based or egg-based additives and thus are vegetarian but not vegan. Similar to the meat industry, egg, and dairy production methods are major sources of animal welfare concerns. In the egg industry, millions of male chicks that are not suitable for egg production or meat production, are killed each year and the beaks of female birds are trimmed to prevent pecking 77 . In the dairy industry, dairy cows are repeatedly impregnated for continuous milk production and are routinely separated from their calves, which are transported to other farms for veal production; causing extreme emotional distress 78 . Even vegan PBM can have indirect effects on wild animal welfare in the form of habitat destruction. To meet food demands, natural vegetation is cleared with monocultural crops which impacts biodiversity. In 1994, palm oil cultivation in Malaysia was found responsible for decreasing mammal inhabitants from 75 to 10 species per hectare 79 . While all agribusiness has an impact on animal welfare that are worthy of concern, substitution of ABM with PBM is still a substantial improvement for animal welfare, as it avoids the unethical treatment of animals during rearing, transportation and slaughter 80 .

One of the primary proposed benefits of CBM is the improvement of farm animal welfare via supplanting intensive animal agriculture. Animal donors are used to supply initial sources of cells that are subsequently expanded in vitro, without needing further resources from the animal. Donor animals, usually younger animals that have more proliferative cells, are anesthetized by a veterinarian and a small (<1 g) tissue biopsy is removed. Cells could be genetically immortalized to proliferate indefinitely; eradicating the need for animal donors. However, in practice, animal donors will likely be relied upon to maintain genetic diversity and to supply non-genetically modified options for CBM. Aside from cell sourcing, a key aspect of CBM production that is linked to animals is serum supplementation. Fetal bovine serum is a common additive to cell culture media and it provides essential growth factors for mammalian cell culture. In 2003, it was estimated that blood from 1,000,000 bovine fetuses was harvested to generate the annual production of 500,000 liters of fetal bovine serum 81 . Serum-free alternatives include co-culture approaches or supplementation with recombinant growth factors.

Looking forward

Animal-based meat production has evolved over thousands of years to supply the demand for affordable and appetizing food. Unfortunately, this feat is accompanied with unintended consequences for human health, natural resources, and the animals involved. Driven by both the rise in global meat demand and increased concern about the aforementioned negative externalities, researchers, and entrepreneurs are turning their focus towards animal-free approaches of meat production (Box 1 ). The economic opportunity for meat alternatives is large and there is no need to crown a single front-runner technology to monopolize the market. Instead, it is important to pursue multiple solutions simultaneously to provide a range of products to serve disparate segments of the consumer market. Plant-based and cell-based meat technologies have made significant advances since their conception. PBM has evolved from being a lackluster meat alternative, that provides a nutritional but not sensory replacement to meat, to be a novel analog nearly indistinguishable from the ABM it seeks to emulate. Likewise, CBM has matured from being the musing of science fiction to being a tangible prototype.

PBM products lie on a spectrum where one end houses more “natural,” less-processed proteins that fall in line with “clean-label” viewpoints but do not do well to mirror the experience of eating meat, while the other contains sensory equivalents that require source proteins to be entirely transformed, and thus viewed as highly processed, and which may come at the cost of certain nutritional factors. Mycelium-based meat may be an exception, where biofermentation can be employed to utilize natural structures and growth patterns of filamentous fungi to mimic meat structures. Along these lines, screening new protein sources that exist in nature that may emulate meat without excessive human manipulation may be an approach that appeals to a wider pool of consumers. CBM is impeded by high production costs, scale-up hurdles, and gaps in fundamental knowledge surrounding how to employ cell culture for food applications. In particular, there are no peer-reviewed, comprehensive datasets detailing the cost, sensory properties, or nutritional value of cell-cultured tissues. While published assessments of environmental impact projections exist, they are based on theoretical large-scale processes that have yet to be validated by industry. To that point, it will be necessary to ascertain details, such as the parameters of material inputs (e.g., doubling time, maximum cell density, medium composition) and industrial-scale production schematics (e.g., bioreactor design, operations) before cost, impact and food safety can be reliably analyzed.

There are opportunities for plant-based and cell-based hybrid products. Considering the current high-cost hurdles associated with CBM, one approach is to focus on the aspects of PBM that fall short of ABM and determine where CBM can add the most value at the lowest inclusion rate. For example, combining PBM with cell-cultured fat may improve the sensory properties of the analog while remaining less costly than a pure CBM product. To this end, expanding research goals to answer fundamental questions surrounding the cost, sensory and nutrition profiles is important to further inform stakeholders on the best areas of application for CBM. Finally, eventual success of PBM and CBM in the marketplace could transform, rather than eliminate, ABM production. If demand for lower quality, previously factory-farmed meat can be supplied with PBM and CBM, demand for higher quality, ABM could be met by smaller-scale, more sustainable and more humane methods of animal farming.

Box 1. A summary of key comparisons between animal-based, plant and fungi-based and cell-based approaches to meat production.

History & Approach Meat consumption has been commonplace since the beginning of human evolution, while plant-based meat analogs are relatively recent dietary additions. Cell-based meat products have not been commercialized, but people have taste-tested prototypes produced by companies and via a singular academic study. Approaches to meat production can be primarily differentiated by starting material: animals (ABM), plants or fungi (PBM) or cells (CBM).

Economics Advances in farming technology and intensification of animal agriculture have resulted in inexpensive and accessible ABM. Although raw inputs for PBM (i.e., unprocessed plant proteins) are less costly than for ABM (i.e., livestock), PBM retail prices are consistently higher due to costs associated with post-processing, production scale and supply chains. Current CBM production is not commercially feasible. Increases in production scale and culture medium-cost reductions are necessary to improve the economics of the process.

Regulatory framework Regulation of novel foods such as certain PBM products and all CBM products is a current topic of interest. Namely, there is a lack of precedent for oversight of food containing cultured cells. Novel additives for PBM may fall within existing regulatory frameworks (e.g., Novel Food Regulation (Europe), “Generally Recognized as Safe” (United States) pathways). There are ongoing debates surrounding labeling laws of what products can be characterized as meat.

Organoleptic properties The primary sensory properties of meat are appearance, aroma, flavor, and texture. Next-generation PBM products are increasingly successful at mimicking processed ABM products (e.g., burgers, deli meat). Current strategies for producing structured PBM products (e.g., filets, steaks) include extrusion, shear cell technology, 3D printing, and mycelium cultivation. Early CBM prototypes appear to emulate ABM but few people have been able to taste-test these samples. The first published sensory panel feedback of a CBM prototype (i.e., cow cells on a texturized soy protein scaffold) reported “a pleasant meaty flavor” and “a typical meat bite and texture”.

Nutrition ABM is a good source of essential amino acids, minerals and vitamins. Some nutritional benefits of traditional PBMs include an absence of cholesterol while providing sources of dietary fiber and healthy fatty acids. Improvements in organoleptic properties may come at the cost of certain nutritional aspects. For example, notable novel PBM products contain high sodium content. There are no publicly available datasets regarding the nutritional profile of CBM. Proponents of the technology assert that nutrition can be regulated by adjusting culture medium formulations and implementing co-culture strategies or genetic modifications.

Consumer acceptance The motivations behind consumption of PBM vary depending on consumer nationality and China and India report higher consumer acceptance rates of PBM compared to the United States. Next-generation PBM products can be perceived as “highly processed” and may not be appealing to “clean label” consumers. With regard to CBM, plant-based consumers are more likely to agree with CBM’s proposed benefits but are less willing to try it compared to omnivores. Consumer perception of CBM is influenced by marketing focus (e.g., process vs. product vs. impact) and terminology (e.g., lab-grown vs. clean vs. cultured).

Public health Overconsumption of red and processed meat is linked to a number of health concerns. There is some evidence that substituting PBM analogs for ABM products can decrease risk factors associated with cardiovascular disease. Compared to ABM, the production of PBM is less associated with pathogenic disease and antibiotic resistance issues. The impact of CBM on public health issues depends on how large-scale production schemes evolve (e.g., sterile processes, antibiotic use). Some consumers report concerns about the safety of CBM products, citing apprehension about unnaturalness and inadequate regulations.

Environmental sustainability Life cycle analyses conclude PBM products are generally more environmentally sustainable than animal-based beef. Some metrics (e.g., greenhouse gas emissions, energy consumption) for some novel PBM products are less sustainable when compared to animal-based poultry. PBM water footprints vary widely depending on the main protein source. The environmental impact of CBM is highly debated, with preliminary assessments presenting significant degrees of uncertainty; especially for energy consumption values.

Animal Welfare Billions of animals suffer and die each year as a direct result of ABM production. Vegan PBM products do not have as direct impacts on animal welfare but crop cultivation can contribute to the destruction of wildlife habitats. One of the primary proposed benefits of CBM is the improvement of farm animal welfare. However, CBM production currently utilizes donor animals for cell acquisition and the culture medium is composed of animal-derived components. The creation of immortalized cell lines and animal-free culture medium ingredients are proposed to address these issues.

Acknowledgements

We thank our lab team members (Andrew, Kyle, John) for stimulating and productive discussions. We thank New Harvest, the Good Food Institute, the Advanced Research Projects Agency, and the National Institutes of Health (P41EB027062) for support.

Author contributions

D.K. and N.R. envisioned the topic and scope of the review. N.R. took the lead in writing the manuscript and figure construction. N.X. aided in writing review sections concerning plant-based meat and participated in figure construction. D.K. and N.X. provided input on all sections of the review.

Competing interests

The authors declare no competing interests.

Peer review information Nature Communications thanks the anonymous reviewer(s) for their contribution to the peer review of this work.

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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Plant-Based Meat Analogues from Alternative Protein: A Systematic Literature Review

Izalin zahari, karolina östbring, jeanette k purhagen, marilyn rayner.

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Correspondence: [email protected]

Received 2022 Aug 26; Accepted 2022 Sep 13; Collection date 2022 Sep.

Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license ( https://creativecommons.org/licenses/by/4.0/ ).

This study aimed to conduct a systematic literature review (SLR) of the research performed in the plant-based meat analogues area. Historical, current, and future tendencies are discussed. The paper offers a comprehensive SLR coupled with a bibliometric analysis of the publication from 1972 to January 2022. The articles were obtained using a research string and precise inclusion and exclusion criteria from two prominent databases, Scopus and Web of Science (WoS). The Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) flow technique was used to describe the data screening and selection. In total, 84 publications were selected for further analysis after a thorough literature assessment. From this study, six main themes were identified: (1) objectives of the study; (2) type of plant protein; (3) product type; (4) added ingredients; (5) texturization technique; and (6) quality assessment considered in the studies. Recent trends in publication imply that meat analogue technology is gaining prominence. This review revealed significant research on improving meat analogues via texturization. Even though extrusion is used industrially, the technique is still in its infancy and needs improvement. Future studies should focus more on fiber and protein–protein interactions, macromolecule conformation and mechanisms, diversifying or improving current methods, sensory attributes, and gastrointestinal absorption rate of each novel protein ingredient.

Keywords: systematic literature review (SLR), bibliometric analysis, meat analogues, meat substitutes, high-moisture meat analogues (HMMA), texturized vegetable protein (TVP), plant-based protein, alternative proteins, texturizing technique, extrusion cooking

1. Introduction

Meat is a major source of dietary protein. It is frequently recognized as a high-quality protein source due to its nutritional qualities and favorable sensory properties such as texture and flavor. However, a rising global population has led to a rise in the production and consumption of meat around the world [ 1 ], which has raised environmental concerns regarding the usage of land and water, as well as the impact of pollution and climate change, greenhouse gas emissions, and the loss of biodiversity [ 2 ]. The inefficient resource usage per gram of protein consumed and greenhouse gas emissions from animal farming have led to recommendations to minimize meat consumption. As the number of European vegetarians, vegans, and flexitarians limiting their meat consumption has increased over the few last years [ 3 , 4 , 5 ], the market for plant-based meat is expected to reach a key milestone of $30.9 billion by 2026 [ 6 ]. In addition, religious concerns and expensive production costs [ 7 , 8 ] also contributes to the transition from animal protein to non-animal protein diets. Plant proteins seem to be a possible solution to these issues since they can replace meat through the creation of nutritionally and structurally equivalent meat-like products. These products are referred to as meat substitutes. Some terminology for meat substitutes includes meat replacers, meat analogues, meat imitations, nonmeat protein alternatives, meatless meats, man-made meats, artificial meats, meat-like meats, mock meats, faux meats, and fake meats. These can be partial or full substitutes for meat, and there is an extensive range of textures. The term “meat analogue” usually refers to products that have a similar look, texture, taste, and color to meat but do not include any meat [ 9 , 10 ].

Over the years, the application of several protein texturizing techniques has been extensively investigated in order to produce meat analogues or meat substitutes. The concept was first developed in the 1970s in an attempt to develop a texturizing technique by using several types of proteins, mainly soy. Isolated soy protein was added to defatted brewers’ yeast to boost protein content and improve the texture of meatball, wiener, and hamburger formulations [ 9 ]. This idea has led to the development of several research areas. Among them are studies conducted by Taranto [ 11 ], who evaluated the function of the extrusion screw in texturized soy and glandless cottonseed flour, and Byun [ 12 ], who showed that soy protein isolates admixtures can be spun using lab-designed equipment. This initial endeavor resulted in several types of resembling meat, so-called meat analogues (moisture content of 50–80%) and texturized vegetable protein (moisture content below 30%). TVP is often combined with real meat to stimulate the texture and appearance of real meat. While HMMA has a fibrous structure, it may be used as a whole meat substitute, which is gaining popularity in the food industry [ 13 ].

In manufacturing HMMA utilizing extrusion technology, protein is the most crucial element. Because of the mix, shear, heat, pressure, and cooling ( Figure 1 ) obtained in the extruder and cooling die, the protein is denatured, unfolded, realigned, and cross-linked during extrusion ( Figure 2 ) [ 14 ]. Protein interactions are regulated by additional factors, such as protein type, pre-treatment, and extrusion parameters, in addition to the types of protein bonds present in HMMA [ 15 ]. Past studies have shown that shearing a protein dispersion in a cone-shaped shear cell may create finely fibrous structures that mimic flesh. Couette cells ( Figure 3 ) with a rotating inner cylinder replaced shear cells (batch systems) for semi-continuous operations [ 16 ]. High-temperature shearing of protein mixtures in a wide-gap Couette shear cell has also led to anisotropic, meat-like structures. Rotation causes a simple shear flow in the protein dispersion, which aligns the proteins and causes solidification. Compared to extrusion cooking which is a continuous process and is faster, the shear cell technique in a batch process may create large, thick, fibrous particles but requires at least 20 min of residence time, while the Couette cell can be operated in a continuous mode and is easy to handle [ 17 ]. On the other hand, the spinning technique is a complicated process that spins high-concentration plant protein into thin fibers that mimic meat ( Figure 4 ). The high voltage between the nozzle and the grounded collector drives electrospinning. In addition, this delicate and costly method uses acid/alkaline solvents and water, causing considerable waste. Electro-spinning mixes protein with other polymers based on solubility, viscosity, conductivity, and other factors [ 18 ].

Illustration of high-moisture extrusion (HME) with twin-screw extruder adapted from Zahari et al. [ 14 ].

Changes in protein conformation during extrusion of HMMA. Modified illustration adapted from Zhang et al. [ 15 ].

Illustration of conical shear and Couette shear cell adapted from Dekkers [ 16 ].

Illustration of electrospinning process adapted from Nieuwland et al. [ 18 ].

In terms of plant protein sources, soybeans were widely employed in the beginning (the 1970s) as a meat substitute such as tofu and tempeh, while wheat gluten was renowned for seitan [ 19 ]. It was reported that in 1976, a total of about 38,700 metric tons (MT) of texturized soy protein (TSP) was produced, which was equivalent to 14% of the total amount of soy flour produced for human consumption [ 20 ]. According to Taranto et al. [ 11 ], McAnelly (1964) textured defatted soybean flour by pre-moistening the flour before exposing it to steam and pressure heating. Texturized protein products were then created by Taranto using extrusion and non-extrusion techniques from defatted soy and glandless cottonseed flours. To this day, researchers have focused their efforts on generating the most palatable meat substitutes from alternative protein sources other than soybeans. These meat substitutes may be created using alternative protein on their own, or with soy protein or wheat gluten added to the mix. Additionally, there has been an increase in the number of studies examining not only the various types of protein materials and texturizing techniques, but also the internal mechanisms, chemical and physical properties, nutritional values, protein interactions, and fiber formation that underpin the mechanism of the products produced [ 21 , 22 , 23 , 24 , 25 , 26 , 27 ].

Various review articles have been published on meat analogues and TVP processing from different angles. In terms of extrusion, Zhang and colleagues [ 28 ] reviewed 115 articles on TVP production, which focused on the effects of barrel temperature, moisture content, feed rate, and screw speed on TVP quality. Other extensive discussions on extrusions have been done by several authors, which covered the aspects related to raw materials, functional properties, and physicochemical changes during the process [ 29 , 30 , 31 , 32 ]. Several techniques used to make fibrous products that mimic muscle meats were discussed by Dekkers et al. [ 33 ]. Furthermore, there has been a long debate about the molecular, nutritional, and functional properties of alternative protein sources as compared to meat [ 34 ]. Some reviews focused on the ingredients used in meat analogues including plant by-products [ 35 , 36 ] and also compared them with traditional meat products [ 37 , 38 ]. An interesting study that reviewed advances in the physical functionality of proteins from non-animal sources in the past three years concerning their potential as meat analogues was carried out by Owens and Levary [ 38 ]. The reviews in recent times have focused on more specific aspects of meat analogues, such as consumer acceptance [ 39 ], structure design [ 10 ], additive ingredients applied in meat analogues [ 40 ], and methods for testing the quality of meat analogues [ 41 , 42 ].

This study aimed to conduct a systematic literature review (SLR) of the research conducted in the plant-based meat analogues area. The purpose of this review was to provide a comprehensive overview of the rapid growth of emerging protein texturization technologies, which need periodic evaluations to keep researchers up-to-date. Our contribution of this paper to the scientific literature is a thorough, up-to-date assessment of the evolution of a large variety of plant-based proteins. The comprehensive literature review was analyzed using bibliometric networks to investigate the relationship between authors and content. Thus, a literature review on meat analogues was conducted to answer the following research questions:

To what extent has research been conducted on the development of meat analogues derived from plant protein-based materials?

What are the key research themes in the literature on plant-based meat analogues?

This review may assist in linking information across several main elements and also in introducing new ideas and encouraging innovation. This paper is divided into the following five sections: An introduction is provided in Section 1 , and the research methodology is described in Section 2 . The findings and outcomes for the first question (RQ1) investigated in this article are presented in Section 3.1 , Section 3.2 and Section 3.3 . The second research question (RQ2) is addressed in Section 3.4 . Some recommendations for future studies are suggested in Section 4 , and the conclusion is provided in Section 5 .

2. Methodology

2.1. overview of review protocol.

A systematic literature review (SLR) is one way to perform a more complete study of the present body of information. SLR aims to discover and synthesize all relevant research in an organized, open, and reproducible manner at each stage of the process. Its ultimate goal is to conduct a full search and analysis of the relevant research [ 43 ]. A comprehensive approach known as PRISMA (Preferred Reporting Items for Systematic Reviews and Meta-Analyses) was used to conduct a systematic literature review that specifies the process of study selection and rejection ( Figure 5 ). This method has been effectively used in a variety of academic fields and can rapidly synthesize key discoveries from an existing knowledge base [ 44 ].

Flowchart of the systematic review search process based on PRISMA guidelines set out in Moher et al. [ 45 ].

2.2. Literature Retrieval and Selection

To answer the research questions (RQs), the advanced retrieval functions in the Scopus and Web of Science (FSTA resources) collection databases were used to retrieve the relevant papers according to the criteria. A search string ( Table 1 ) was developed to ensure the quality of the literature. This study was performed in 2019 and was periodically updated until January 2022. The preliminary search yielded 1603 records. For inclusion criteria, the document categories were firstly limited to peer-reviewed scientific journals and English-language articles, excluding forms such as conference papers, proceedings, book chapters, reports, and editorial materials. A total of 851 records was left after this first-round inspection. Next, data from accepted articles were loaded into Endnote (online bibliographic management software), version basic, Clarivate Analytics ( https://access.clarivate.com/#/login?app=endnote , accessed on 1 February 2022). Endnote software facilitates duplicate removal. After removing the duplicates (61 articles), the remaining articles was retrieved and evaluated for relevance by carefully reading the title, keywords, and abstract.

Search string used in the selected database.

* Truncation of terms was used to capture variation in language.

The review’s scope included 227 articles, and the content was evaluated by reading the full article (abstract, method, results, and conclusion). Following that, 143 articles were eliminated as undoubtedly out of the review’s scope. Review articles and those without the full text were removed. At this stage, there were still several articles written in languages other than English (the titles were the only part that was translated), and duplicates that needed to be eliminated. The inclusion and exclusion criteria for this round focused on whether the document was consistent or not with the research questions, which were: (i) studies focused on textured vegetable protein (TVP) or meat analogues processing from plant-based protein (non-animal based which include fungi, algae, and mushroom); (ii) studies focused on quality assessment of protein materials or product developed or both. In particular, the following topics were excluded: (i) non-TVP or non-meat analogues products such as expanded products (snacks, cereals, and pasta); (ii) studies involving animal-based products (meat, milk, fish, blood, whey) or insects based raw materials added in TVP or meat analogues; (iii) other textured products such as 3D printed products, cell-cultures products, food ingredients, flour, cookies, bakeries, pet food, drugs, and non-food products; and (iv) unrelated subjects such as studies mainly on consumer acceptance, sensorial techniques, economic aspects, or marketing (without the texturization process). To be clearer, studies were only selected based on plant protein materials, processing into texturized protein, and/or meat analogues. There were also some articles that were discovered to be totally unrelated only after reading the full articles. In total, 767 records were removed, leaving 84 full-length articles in the review portfolio.

2.3. Bibliometric Analysis

To investigate the chosen relevant publications, bibliometric techniques such as co-authorship analysis and keyword co-occurrence analysis were used to trace the knowledge architecture of a particular study subject. It is also a technique for detecting the patterns in development or the directions in which future research should go [ 46 ]. These are the most thorough procedures utilized to illustrate the relationship between the author and the keyword cluster used in the selected articles. VOS viewer software was used to perform the clustering. The VOS acronym stands for “visualisation of similarities”, and the program is an open-source application ( www.vosviewer.com accessed on 1 February 2022) developed to create and visualize bibliometric maps [ 47 ]. It has been shown that network visualization is a valuable tool for analyzing a wide range of bibliometric networks, such as networks of co-authorship relationships among academics, and networks of keyword co-occurrence associations in published works [ 48 ].

3. Results and Discussion

3.1. publication trends.

Figure 6 illustrates the number of articles based on the publication year, which varied across the years. The first publication on the texturization of plant protein for meat substitutes appeared in 1972, and the findings revealed that there has been a substantial quantity of scientific production since then. It particularly peaked in 2021 when 18 papers on the topic were published, a trend that may continue in the coming years ahead, considering that the database search for 2022 only included papers available until January.

Annual publication trend of 84 papers between 1972 and January 2022 retrieved from Scopus and Web of Science (WoS).

3.2. Journal-Based Publications

Table 2 presents the list of the most prominent journals (top 13) publishing articles on meat analogues or TVP and development in the field. The bibliometric analysis revealed that the 84 papers included in the systematic review were published in 38 journals between 1972 and 2022. It was revealed that the most frequent journals chosen by the authors were Foods with 15 articles [ 14 , 15 , 24 , 41 , 49 , 50 , 51 , 52 , 53 , 54 , 55 , 56 , 57 , 58 , 59 ], followed by Journal of Food Science with nine articles [ 11 , 20 , 25 , 60 , 61 , 62 , 63 , 64 , 65 ]. Because the journals were indexed by both Scopus and Web of Science databases, they were all of high quality, and most were placed in Quartile 1 or 2 (see Table 2 ).

Journals with the largest number of documents.

3.3. Meat Analogue Bibliometric Networks

3.3.1. network visualization for author keywords.

In the network presented in Figure 7 , each circle represents a keyword. A node depicts a term in a publication title or abstract. The size of a node indicates the number of publications in which it is present. The nodes are colored based on this journal’s average year of occurrence using the color scheme depicted in the legend. Nodes closer to each other are more similar than nodes further apart. The thickness of a link between two nodes indicates the likelihood that they co-occur in the same publication. The minimum number of occurrences of the keywords is three. From 594 keywords, 71 meet the threshold. It was found that a number of authors utilized variants of keywords. The most frequently occurring terms were “extrusion” (27 documents), “protein” (25 documents), and “meat” (16 documents), followed by the terms “moisture” (15 documents), “meat analog” (14 documents), “plant protein” (13 documents), “meat analogues” (11 documents), and “meat analogue” (10 documents). Early in the twenty-first century, there was an increased interest in “soy proteins”, “glycine max (soy bean)”, and “triticum aestivum (wheat)”, as well as “flour”, “vegetable protein”, “gluten”, and “plant protein” (average publication year: 2005–2010). In recent years, authors have increasingly focused on “extrusion”, “meat analogues”, “high moisture extrusion cooking”, “functional properties”, and “mechanical properties” (average publication year: 2015–2020).

Overlay visualization based on keyword re-occurrence of at least three times based on 84 retrieved articles.

3.3.2. Network Visualization by Author

Our search revealed that 297 authors contributed to the total number of publications based on the selected 84 articles. Figure 8 depicts a network visualization of the authors’ co-occurrence network for all the publications analyzed in this search, which formed 56 clusters. Each circle indicates a different author’s name. The size of the circle is proportional to the number of articles each author has written on this subject. In general, the greater the proximity of authors in the visualization, the stronger their bibliographic connection. Only 43 authors had two or more publications, whereas 254 authors had only one publication. Emin, M.A.; Hsieh, F.; and Kabstein, H.P. had the most publications in this dataset with five each, followed by Yao, G. with four publications. In addition, the co-authorship network’s largest connected subgraph included just 18 authors, including Van der Goot, A.J.; Krintiras, G.A.; Cornet, S.H.V.; and Jia, W. The second important subgraph in the co-authorship network consisted of 15 authors, including Grahl, S.; Palanisamy, M.; Saerens, W.; and their peers.

Network visualization based on authors in 84 retrieved articles.

3.4. Classifications

One of the most important things to report on in SLRs is the table of findings. The table clearly explains the results and makes them easy to understand [ 44 ]. Table 3 shows all selected articles, which were classified according to the content, resulting in six main themes: (1) objectives of the study; (2) type of plant protein; (3) product type; (4) added ingredient; (5) texturization technique used; and (6) quality assessment considered in the studies. These six themes further produced 22 sub-themes, which provided answers to the second research question (RQ2) of this SLR. The background of the selected studies is discussed in the following section.

Findings from 84 reviewed articles.

3.4.1. Objectives of the Reviewed Studies

The articles chosen were well-balanced in terms of their objectives. Table 3 demonstrates that the majority of the included articles studied the texturization technique in creating texturized vegetable protein (TVP) or meat analogues (MA), as well as protein materials using either commercial soy protein or other protein-based materials. Some studies investigated how fiber development evolved in the MA/TVP. Another critical aspect of meat substitutes is quality assessment, which was extensively identified in many articles. We proposed a framework to address three main aspects highlighted in the reviewed articles, as seen in Figure 9 .

Overall framework composed of three components.

3.4.2. Type of Plant Proteins Used

Soy protein as primary component.

In this first sub-theme, it was discovered that experts had focused their attention on soy protein (either in the form of flour, concentrate, or isolate) as the primary component in textured protein products for years. Soy protein is used either by mixing it directly in the formulation of meat substitutes together with other ingredients, or by processing it via texturizing techniques into TVP or HMMA. Soy garnered interest for its protein quality and because it has satisfactory functional qualities (such as the ability to absorb water and oil and its emulsifying properties), and it has been used in the production of a variety of unique meat substitutes. As a result of the excellent features that soy has, it is usually used as a standard or benchmark to compare different protein materials [ 78 ] and as a model to explore many other aspects of meat analogues [ 63 ], texturizing techniques [ 19 ], extrusion parameters [ 15 ], and product structure [ 58 , 59 , 73 ]. Dahl and Villota [ 60 ] used soy flour altered with acid (HCl) or base (NaOH) and studied the pH effects on the functional properties of soy protein. Liu and Hsieh [ 88 ] used two commercial soy protein isolates to study the fibrous meat analogues produced through high moisture extrusion or gels via heating and chilling, with different concentrations and/or temperatures. Due to its lower cost, soy protein concentrates (SPCs) are widely utilized as an alternative to soy protein isolate (SPI). Pietsch et al. [ 100 ] reported that SPCs may produce more prominent anisotropic (properties of materials depending on the direction) structures than SPI. Two other studies using high-moisture extrusion of soy meat analogues (SPCs) were conducted by Palanisamy et al. [ 98 ] and Chiang et al. [ 72 ].

Soy Protein Combined with Other Plant Proteins

Researchers started to use commercial soy protein isolates/concentrate together with other protein sources in order to reduce the use of soy protein, and also to study their combination, establish texturization conditions, and to aim to diversify meat products in the market with different formulations, as discovered by many authors [ 15 , 61 , 62 , 70 ]. Kozlowska et al. [ 81 ] used high- and low-pressure processing to texturize the flours and concentrates that were derived from soybean and rapeseed, as well as the blends of soybean and rapeseed (1:1). When combined with additional plant proteins, the extruded meat analogues were found to be of higher quality. Numerous researchers have found that by combining soy protein and wheat gluten, meat substitutes might match the texture, color, flavor, and function of red meat, as well as enhance the disulphide bonds to generate a fibrous structure [ 27 , 72 , 111 ]. In the investigation of the total heat transfer coefficient in extrusion processing conducted by Lee et al. [ 84 ], meat analogues were mixed with another established protein and wheat starch. In a separate investigation, Liu and Hsieh [ 89 ] and Ranasinghesagara et al. [ 25 ] similarly co-extruded soy protein with wheat gluten and starch to produce fibrous meat analogues under high-moisture and high-temperature conditions. According to prior studies by Neumann et al. [ 95 ], non-heated corn gluten (CG) demonstrated superior functional performance compared to heat-dried corn gluten meal. Thus, wet-milled corn gluten and defatted soy flour (DSF) were combined and extruded to produce textured meals afterwards. Hemp protein could also be mixed with soy protein isolate up to 60% in the formulation of high moisture meat analogues, as reported in a previous study [ 15 ]. In the context of restructured meat analogues, mushrooms, which contain high levels of sulphur-containing amino acids, and glutamic acid, which implies a distinctive umami taste, are the materials that are mostly used, as they closely resemble those with a natural meaty flavor and texture. Because of these similarities, mushrooms have been employed by mixing directly with other added ingredients in the formulation of several types of restructured meat analogues [ 66 , 77 , 80 , 94 ]. Other plant proteins, which were recently found to be promising to partially substitute soy protein in meat analogues, include spirulina [ 76 ], yam [ 87 ], rice protein isolate [ 85 ], and microalgae powder [ 71 ].

Alternative Proteins without Soy Protein

The second sub-theme pertained to alternative plant proteins used in previous studies. Researchers examined other protein resources to completely replace soy protein in the formulations. This interest is due to other factors, including GMO issues, allergies, and an unfavorable climate for soy cultivation. However, in thermomechanical processing that involves texturizing equipment, it is impossible to make great meat analogues without the use of components that have a high percentage of protein. To develop a comprehensive fibrous structure similar to actual tissue, extrusion, shear cell, and spinning technologies, for example, need ingredients with a high protein concentration. Several studies showed promising protein ingredients such as pea protein [ 31 , 93 ], mucuna beans [ 96 ], peanut protein [ 103 , 114 ], and faba beans [ 24 , 105 ]. The majority of the plant-based proteins (such as those found in legumes and oilseeds, for example) contain undesirable components such as anti-nutrients (glucosinolates, phenolic compounds, and phytic acid) and inhibitors of digestive enzymes. These components reduce the nutritional value and acceptability of plant-based proteins and impart an unpleasant flavor such as bitterness. To be accepted by consumers as meat analogues, these undesirable components must be eliminated by certain pre-treatments before being used. Those treatments used together with the protein extraction process, on the other hand, lead to a loss of functional qualities as well as reduction of the protein’s quality and quantity. This is by far the most challenging obstacle to overcome when researching and developing novel plant protein materials. Because various plants have varying protein types and qualities, numerous efforts have been made to create a novel blend of several plant proteins with the hope that some proteins may compensate for the drawbacks of other plant proteins during the texturization process. For example, Kozlowska et al. [ 81 ] and Zahari et al. [ 14 ] suggested that rapeseed protein would be a good source for supplementing other vegetable proteins, e.g., soybean and yellow pea. Arueya et al. [ 67 ] created meat analogues from Lima bean protein concentrate (LBPC) and African oil bean seed concentrate (AOBSPC), which are underutilized legumes with high nutritional potential grown mainly in Peru. Similarly, De Angelis et al. [ 49 ] successfully developed meat analogues by employing different protein mixtures from dry fractionated pea and oat protein. In summary, meat analogues produced with other plant proteins than soy have a distinct fibrous structure, high levels of vital amino acids, and a nutritionally useful composition, making them prospective future elements.

3.4.3. Product Type

Two sub-themes were developed under this theme: texturized vegetable protein (TVP), and meat analogues (MA). Regarding nomenclature, there is a considerable degree of disagreement among professionals, with some arguing that extrudates from extruders cannot be referred to be meat analogues, while others disagree. Some authors suggest that the extrudates are not intended for immediate consumption but rather as meat extenders that will be sliced and combined with other substances to form a restructured meat substitute. Thus, depending on the publication, some of the studied articles referred to the extruded product as extrudates or meat analogues, while others referred to it as TVP and utilized it as a replacement for meat. We categorized the terms based on the information provided in the articles, including the final restructured meat-substitute products, which are normally referred to as nuggets, sausages, or patties and may contain added ingredients according to the formulations.

Texturized Vegetable Protein (TVP)

In 1978, Hashizume [ 19 ] studied a traditional method of manufacturing Koritofu to convert protein into a textured product using a freezing method. In comparison to the other temperatures that were evaluated in the research, such as −5 and −70 °C, it was claimed that the temperature of −20 °C was the one that successfully created the spongy protein that could be utilized as a replacement for animal flesh. According to Kozlowska et al. [ 81 ], who used two different models of extruder, the high-pressure technique produces a product with a specialized purpose as a meat extender, whereas the low-pressure technique produces a product that is suitable for developing meat analogues. However, Neumann et al. [ 95 ] defined the product as TVP when it was produced by low-pressure extrusion at pressures around 100–200 psi. On the other hand, according to Maung et al. [ 92 ], TVP is often used as a meat extender or directly as meat analogues in hamburger patties, sausages, steak, sliced meats, and many other products. Bakhsh et al. [ 41 ] recently revealed that TVP, when used as a major ingredient in hamburgers, had characteristics comparable to those of hamburgers made from beef and pork. However, it was noted in his other study that the surface of the patties that were made from TVP and texturized SPI both had a granular look, which is a downside of employing those two ingredients [ 68 ]. While using chickpea flour and TVP, Sharima-Abdullah et al. [ 107 ] produced an imitation of chicken nuggets, which was stated to be a promising product.

Meat Analogues

In terms of meat analogues, several researchers developed the products directly as whole muscle meat (mostly from extrusion) with or without some added ingredients [ 24 , 49 , 51 , 52 ], but others referred to meat analogues as restructured meat products such as Turkish dry fermented sausages (“sucuk”) from wheat bulgur [ 69 ], SPI sausage [ 79 ], and edible mushroom sausages [ 57 , 109 ]. The study by Rousta et al. [ 56 ], who investigated the culture of the fungus on oat flour and its use in the development of burger patties, demonstrated the productive potential of the fungus for the manufacture of nutrient-dense foods. Saldanha do Carmo and colleagues [ 105 ] used response surface methodology (RSM) to optimize the manufacturing of meat analogues made entirely of faba bean protein concentrate acquired by a dry-fractionation technique, which also showed promising results.

3.4.4. Added Ingredient Used to Improve Texturized Products

Binding agents.

When making textured vegetable protein products or meat substitutes, it is a common practice to use some amount of additives or chemicals in order to expand the range of raw materials suitable for use in production [ 84 ]. Many different binding agents have been utilized as fat replacers to increase the quality of TVP or restructured meat (for example, sausage, nugget, and patty). This has been done to improve the taste, juiciness, mouthfeel, and other sensory qualities. Examples of such additions are starch, fibers, soy and milk proteins, a variety of hydrocolloids, and egg solids. Because the scope of this study is limited to plant-based products, we will not include any materials derived from animal sources; as a result, we will only count a few studies. Arora et al. [ 66 ] investigated the effect of various quantities of binding agents (carrageenan, soy protein concentrate, casein, and xanthan gum) on the qualitative features and nutritional qualities of mushroom-based sausage analogues prepared with 5% saturated fat. Carrageenan (0.8%) had the greatest outcomes in terms of minimizing purge loss, cook loss, and emulsion stability, all of which improved the process output. It was reported that methylcellulose (MC) is an effective binder, particularly for the meat analogues that do not need to be preheated for gel formation, because of its one-of-a-kind thermal gelling ability and emulsifying qualities [ 41 ]. Contradictorily, Sakai et al. [ 26 ] developed an alternative new binding mechanism, since chemicals are used in the production of methylcellulose. The results suggest that the protein–sugar beet pectin crosslink catalyzed by laccase may serve as a binding mechanism for the TVP patties. In addition, microbial transglutaminase (TG) and sodium alginate (AL) are two binding agents that are often employed in food preparation. Each of these binding agents functions in distinct ways for protein binding or gelling systems [ 26 , 69 , 86 ]. The authors suggest that the combination of TG and AL may synergistically affect the eating quality of soy patties, although more improvements are required. While AL has an advantage in the creation of restructured meat because it can create a thermostable and irreversible gel (in the presence of Ca2+), TG has been used as a cold-set binder since it catalyzes covalent bonds between the ε-amino group (a primary amine) of peptide-bound lysine and the γ-carboxamide group of peptide-bound glutamine [ 26 , 69 ].

In order to reduce the saturated fatty acid and cholesterol levels of certain restructured meat substitutes, animal fats were substituted by vegetable oils such as olive oil [ 26 , 69 ], palm oil [ 55 ], canola oil, and coconut oil [ 41 , 68 ]. Depending on the raw materials, oil is used in different amounts to obtain a more meat-like texture and to increase the flavor, juicy quality, tenderness, and several other qualities of MA related to sensorial experience. For example, Mazlan et al. [ 55 ] used 10% palm oil in the soy–mushroom extrusion mixture. Kamani et al. [ 79 ] used 8% oil in SPI-gluten sausage analogues, Bakhsh et al. [ 68 ] used a total of 7.5% of oil in the TVP patty formulation, while Saerens et al. [ 104 ] used fat emulsion from pea protein and rapeseed oil in the soy and pumpkin seed protein-based burger patty formulation. It has been observed that fat has an influence on thermal–mechanical processing as a lubricating agent and helps to accelerate the creation of protein alignment networks. Recent research conducted by Kendler and colleagues [ 53 ] investigated the impact that oil (0–6%) had on the extrusion-relevant parameters and structure-related properties of extruded wheat gluten. According to the results, using oil in the high-moisture extrusion led to a significant change in the process conditions, as well as in the rheological properties and product qualities. The oil concentration and addition point were discovered to have an impact on the size of the oil droplets. The size of the oil droplets became larger as the oil content increased, indicating that the fat droplets were subjected to coalescence. On the other hand, there was a difference in oil droplet size depending on where in the extruder the oil was injected, where injection at the end of the extruder resulted in smaller oil droplets [ 53 ]. The anticipation that oil droplet breakage is improved at greater matrix viscosities was supported by these findings. Some protein materials still contain a high content of oil, such as rapeseed protein concentrate [ 14 ], which could thus enhance the final product characteristics without adding any fat during thermal–mechanical processing. Nevertheless, too much oil may contribute to the lubricating effect (slippery condition) within the barrel, hindering the protein denaturation process.

Other Ingredients

In addition to binders and lipids, there are a few other ingredients that are normally added into the formulation of meat analogues, especially restructured meat. For extruded meat analogues, added ingredients such as polysaccharides, colorants, flavoring, and seasoning were used during the cooking process. Exogenous polysaccharides are one of the key additions often employed in the food industry to increase the functional qualities of food proteins and optimize texture, and they were utilized to study the impact of polysaccharides in the meat analogues of peanut protein [ 115 ]. Since the majority of extruded meat analogues lack flavor and color, adding additional ingredients such as meat flavor powder and red yeast rice is required when incorporating them into restructured meat, as shown in [ 57 ]. Wen et al. [ 110 ] studied the effect of calcium stearyl lactylate (CSL) in extrusion processing and found that CSL has the potential to greatly improve the extrudates’ textural qualities, including the number of fibrils present and the size of their pores. In terms of restructured meat, the taste of patties and sausages was improved by the addition of colorant, sugar and salt, flavoring, seasoning, herbs, and spices such as cumin, cinnamon, pepper, and garlic [ 57 , 68 , 69 , 109 ]. According to Yuliarti et al. [ 113 ], the inclusion of calcium chloride and baking powder in the formulation of pea and wheat protein nugget was to boost the protein’s ability to bind water and to create air cells in the dough, which might improve the fibrous structure. Carotene and anthocyanin are also being added to enhance the vegetarian sausage analogues; nevertheless, it has been noted that their levels drop with storage, thus requiring additional development [ 109 ].

3.4.5. Type of Texturization Technique

Many texturization technique studies have been conducted in recent years, which were classified into five groups: single screw extrusion (SSE), twin screw extrusion (TSE), shear/Couette cell (SC), spinning (S), and mixing/other methods (M/O). For thermal–mechanical processing, high moisture extrusion technology has become a popular method compared to other texturizing methods due to its lower energy consumption, lack of waste to be disposed of, high efficiency, and higher textured product quality [ 14 , 21 , 24 , 49 , 51 , 53 , 54 , 59 , 101 , 112 ]. In the past, the production of TVP, which often has a lower moisture content, was mostly done using single-screw extruders [ 81 , 96 , 99 , 103 ]. A drawback with TVP is that it must first undergo a rehydration process before being included in the meat substitutes’ formulas. Extrusion seems to be becoming increasingly popular since it can be utilized for both low- and high-moisture products. Because of these factors, several experiments were performed to gain an understanding of the relationship between the processing parameters of the extruder and the final product. According to Samard et al. [ 106 ], who examined the effect of extrusion type (low- and high-moisture extrusion cooking) on the physicochemical properties of meat substitutes, the cooling die region of HMMAs is thought to be crucial for the cross-link formation. As we know, for most plant proteins, a certain pre-treatment (acid or alkaline) is typically required during the extraction process in order to receive a higher yield as well as more desirable extrusion outcomes. Furthermore, extrusion was shown to partly break down phytates in a matrix-dependent way, improving the material’s nutritional quality [ 52 ]. In addition to the composition of raw materials, various extrusion parameters (screw configuration, temperature set-up, screw speed, solid and liquid dosing, and moisture content), as well as diverse manufacturers, can result in dramatically different product structures and textures, even when using the same raw material. Each processing parameter will influence the product specification during extrusion processing. For instance, most studies investigated the effect of screw speed, water feed, extrusion temperature, and feed rate on the product characteristics [ 14 , 78 , 100 , 102 , 106 ]. Studies showed that water feed was the most influential factor in the extruder process and product qualities, followed by screw speed and barrel temperature [ 14 , 97 ]. Several blocks of barrels could be used in the high-moisture extruder, with the highest temperature ranging from 100 to 180 °C [ 97 , 100 , 102 , 103 , 106 ]. Earlier, Lee et al. [ 84 ] calculated the total heat transfer coefficient in a long slit cooling die and discovered that the projected product temperature at the die output was 6.8 °C of the observed experimental value, whereas several authors investigated the effects of the specific mechanical energy (SME; kJ/kg) [ 14 , 74 , 100 ] and specific thermal energy (STE; kJ/kg) [ 71 ] on the physicochemical properties of texturized meat analogues. Increasing the screw speed required more energy, but the SME dropped as the moisture content increased [ 14 , 71 ].

Shear/Couette Cell

Several studies have suggested the shear cell as a suitable technique for meat analogues production. A cone–cone device (shear cell) and a concentric cylinder device (Couette cell) were created by Krintiras et al. [ 17 , 82 , 83 ] based on the notion of a flow-driven structure. In both devices, a model system of soy protein isolate (SPI) and vital wheat gluten mix was employed, resulting in anisotropic structures that may be used as meat replacers. In 2016, they invented a 7 L Couette cell system for making structured soy meat replacer, and high anisotropy fibers were developed. The up-scaled Couette cell can produce 30 mm thick flesh replacers that mimic meat, which means that the research found no impediments to scaling up the idea. The flexible design enables the manufacturing of meat substitute goods in sizes not previously possible, which might be beneficial in replacing chicken breast or beef [ 17 ]. According to Jia et al. [ 78 ], the formation of fibrous materials in shear cells is favored when plant materials have two different phases that deform and align when sheared. This can be done by mixing purified ingredients with different water holding capacities, such as soy protein isolate and wheat gluten, or they can be found naturally in a single but less purified ingredient, such as soy protein concentrate. They studied the structuring potential of rapeseed protein concentrate (RPC) with and without wheat gluten (WG) for meat analogues synthesis in a shear cell. Both RPC-only and RPC–WG combinations could become fibrous at 140 °C and 150 °C with 40% dry matter; in addition, WG could enhance the fibrous structure and lighten the color [ 78 ].

In 1972, Stanley et al. [ 108 ] studied the properties and ultrastructure of rehydrated spun soy fiber and identified structural differences between soy and beef. According to the study, meat contains repeated sarcomeres, connective tissue that affects texture, a sarcolemma or elastic cell membrane, and actin–myosin cross-bridges, whereas spun soy is a uniform, homogenous fiber with disulfide bonds. Byun and colleagues [ 12 ] designed bench-scale protein spinning equipment in the laboratory with some modifications to the prior approach in order to determine the viability of spinning mixes of soy protein isolates. The discovered method is based on the unfolding of peptide chains by alkali treatment and the molecular orientation of mechanically spun fiber. Electrospinning is one of the techniques investigated recently by Mattice et al. [ 91 ], and they modified the electrospinning parameters to produce zein fibers with uniform width while minimizing ethanol consumption. Even though electrospinning produces tiny individual fibers, the technology used in this research was reported to have a very low throughput and thus faced problems with efficiency.

Other Texturization Methods

In addition to all of the particular thermomechanical and texturizing procedures, there were also other methods being used in exploring and developing meat analogues, such as direct mixing [ 79 ], freezing [ 19 ], planetary roller extruder [ 98 ], mechanical elongation, and antisolvent precipitation [ 91 ]. According to the findings of this review, the majority of meat analogues that employed mixing techniques with other ingredients was restructured meat products such as nuggets, sausages, and patties [ 77 , 79 , 80 , 87 , 107 , 109 ]. Furthermore, Nayak et al. [ 94 ] developed a meaty-textured soybean by solid-state fermentation using Rhizopus oligosporus and dried Agaricus mushroom and compared the textural profile of the optimized fermented soybean with poultry meat.

3.4.6. Quality Assessment Considered

As previously shown in Figure 7 , improving the qualitative attributes of meat analogues has been the focus of several investigations over the last few decade. These quality efforts are important for developing excellent meat analogues, especially in understanding the formation of the fibrous structure and protein–protein interactions. Meat consumers were less tolerant to plant-based commodities compared to actual meat products a few years ago, and the main reason was the poorer sensory and nutritional value of the plant-based products. Following that, several novel meat substitutes with enhanced flavor and texture from a variety of plant-based sources can be found on the market, the most popular being based on pea and oat. Several elements of meat substitute characteristics, including chemical and functional properties, physical properties, fiber formation, nutritional properties, cooking quality, and sensory evaluation, are being addressed. Here we may observe how meat analogue research contributed to the food industry. Fiber creation has recently been a hot topic in many authors’ research.

Chemical/Functional Properties

The most widely used approach in this sub-theme is proximate analysis. All protein powders and meat substitutes are proximately analyzed using international standard methods (AOAC, AACC, ISO), which include moisture, protein, fat, crude fiber, and ash. In extrusion processing, it is necessary to know the moisture content of the protein materials (feed powder) in order to determine the desired moisture content of the final extrudates. Most of the previous work used a conversion factor of 6.25 for soy protein and 5.7 for wheat gluten [ 31 , 88 , 89 , 114 ], and the protein was examined using either the Kjeldahl or Dumas combustion methods. In the future, additional conversion factors specific to each crop may need to be employed to obtain more accurate findings. For example, Mariotti et al. [ 116 ] proposed a collection of certain conversion factors for various meals, such as 5.5 for soybean, 5.4 for cereals and legumes pulses, and 5.6 for corn and other sources. The study suggested a more exact default conversion factor of 5.6 rather than 6.25, a scientific way to express nitrogen as protein, which is highly relevant when “protein” refers to “amino acids”. Using flame atomic absorption spectrometry, the levels of the microelements such as iron (Fe), zinc (Zn), copper (Cu), and manganese (Mn) in the extrudate samples were analyzed. Many studies investigated protein–protein interactions using the protein solubility approach [ 32 , 88 , 89 , 114 ]. Osen et al. [ 32 ] studied the establishment of covalent peptide bonding during the extrusion process in order to evaluate the impact that high moisture extrusion cooking had on the protein changes that occurred within the extruder. Moreover, FTIR is commonly used for investigating protein conformation and is capable of accurately measuring the secondary structure of proteins, since each protein may be linked with a unique set of bands and wavenumber intensities [ 87 , 110 , 112 , 114 , 115 ]. SDS–PAGE analysis was commonly used to investigate the degree of crosslinking, thus could determine the molecular weight distribution [ 26 , 74 , 101 , 111 ]. Proteins may polymerize into larger aggregates, rendering those proteins too large to penetrate the flowing gel [ 74 ]. As explained by Kaleda et al. [ 52 ], changes in the content and conformation of proteins significantly impact the capacity of proteins to hold water. Water holding is highly dependent on the presence of polar hydrophilic groups, while the nonpolar side chains of proteins are responsible for determining the oil holding capacity of a material. Oil holding capacity also relies on the physical trapping of oil and may be explained by the material’s microstructure. Because it impacts the quality and production of meat analogues, water, and oil holding capacity are important factors. The greater the holding capacity of a product, the juicier it will be. The water solubility index (WSI) measures the total quantity of a substance that can be extracted using water. Multiple variables, including powder composition and particle size, conformational state of proteins, molecular size, and cross-linking, may impact WSI [ 52 ]. Using carrageenan (0.8%) as a binding agent, mushroom sausages exhibited the lowest amount of purge loss (3.56%) after being frozen, which led to a reduction in drip losses caused by the thawing process [ 66 ].

Thermal analysis is usually conducted using differential scanning calorimetry (DSC), thermogravimetric (TG), and differential thermal analysis (DTA) to measure the thermal denaturation of protein, as used in several studies [ 24 , 24 , 88 , 114 ]. This also helps in setting the correct barrel temperature during the extrusion process and increases knowledge of the raw materials. The barrel temperature must be high enough to allow protein denaturation during the extrusion process. A rheology study may provide some insight into the flow and deformation of protein materials. Many studies reported the rheology results previously [ 50 , 60 , 114 ]. It can assess the behavior of proteins under shear stress and strain during heating–cooling cycles and act as a predictor of the quality of the finished product (meat analogues) following thermal–mechanical processing [ 15 , 24 , 100 ]. Emin et al. [ 73 ] employed a closed cavity rheometer (with a specified extrusion-like environment) to investigate the critical process parameters that contribute to a substantial change in the response behavior of a plant protein model system, employing vital wheat gluten as a model system. The findings reveal that temperature, water content, shear, and a step change in shear significantly impact the response behavior of proteins.

Physical Properties

Physical attributes consist of analyses performed on TVP or meat substitutes. Textural and structural properties using a texture profile analyzer (TPA), cutting strength and tensile strength, and ultrastructural characteristics such as scanning electron microscopy (SEM) and light microscopy analysis are among the most typical tests performed to determine how the texture or morphology are formed inside the product and how they are connected to other properties such as chemical composition, SME, and sensory attributes. Authors mostly used SEM to comprehend the structural properties of meat analogues [ 17 , 80 , 81 , 95 , 98 ]. The important characteristics highlighted using TPA were hardness and chewiness. As a result of cutting through the fibrous meat analogues, it was discovered that the values for the longitudinal cutting strength were significantly higher than the values for the transversal cutting strength [ 14 ]. In most cases, the authors either compared the physical characteristics of meat analogues to a reference product of chicken meat and beef or commercial meat analogues. The degree of texturization of SPI-based meat substitutes rises as the SME decreases, according to several studies [ 72 , 74 ]. Kaleda et al. [ 52 ] discovered that this is not the case when employing a screw configuration with several kneading and reverse blocks, resulting in greater mechanical treatment. Lower hardness and chewiness in meat analogues were, on the other hand, reported by Fang et al. [ 74 ], being correlated with lower SME, contrary to Chiang et al. [ 72 ] and Zahari et al. [ 14 ]. In the extruder barrel, protein molecules exhibited major structural changes and unfolding, creating ideal circumstances for molecular rearrangement in the subsequent extruder zones. The meat-like fibrous structure was reported to be formed at the cooling die zone junction due to protein phase separation and rearrangement [ 58 ].

When determining color, the results are always associated with the color of the raw materials used, and the temperatures at which the food was cooked might influence the product. For instance, meat analogues from soy protein had a lighter color compared to hemp, and the L* values decreased as the proportion of hemp powder in the formulation increased [ 15 ]. Some authors agreed that L* and a* values were dependent on extrusion temperature and moisture during extrusion cooking [ 52 , 97 ]. Raising the moisture content would increase the L* values owing to the lower rates of chemical reactions in the protein composite produced with greater water content [ 24 , 97 ]. Other physical properties applied by the authors in this review on TVP and meat analogues are lateral expansion [ 96 ], bulk density [ 49 ], porosity [ 67 ], expansion index [ 60 , 70 ], and rehydration [ 70 ].

Fiber Formation

Yao and colleagues [ 65 ] devised a technique for determining fiber formation in soy protein extrudates with high moisture content. To assess fiber growth, many polarization measurements are required. Because the fluorescence signal is weak, ambient light must be blocked, which is problematic on a production line. Ranasinghesagara and colleagues [ 63 ] refined the technology by establishing an image processing method to automatically quantify fiber creation using digital imaging and by inventing a non-destructive imaging approach with real-time quality control [ 64 ]. Later in 2009, more sophisticated technology was applied by incorporating it into a rapid laser scanning system, which permits real-time 2D mapping of fiber production and orientation across the sample [ 25 ]. Another interesting study was published by Zhang et al. [ 114 ], who employed a multiscale approach paired with emerging techniques such as atomic force microscopy-based infrared spectroscopy and X-ray microscopy to make the entire extrusion process visible to illustrate the process of generating a meat-like fibrous structure. Moreover, simulations show that phase separation under temperature or velocity gradients may lead to multilayer structures [ 93 ].

Nutritional Analysis

In addition to functional and textural qualities, the nutritional composition of meat substitutes is an essential factor to be considered when substituting meat with plant-based analogues. Regarding amino acids, researchers are exploring alternative approaches to meet the FAO’s requirements for meat. Proteins derived from plants are considered nutritionally insufficient because cereals often lack lysine, and legumes typically have low levels of the sulphur-containing essential amino acids methionine and cysteine. As a result, the quality of the nutrients could be increased by mixing two or more protein sources, which enable the mix to meet FAO standards for a particular age group. Several publications [ 22 , 24 , 32 , 67 , 70 ] reported the amino acid composition; however, no publication was found that analyzed the protein quality or protein digestibility-corrected amino acid score (PDCAAS) of the produced meat analogues. This demonstrates that there is currently a lack of information that is readily available. Previously, it was reported that the amino acid composition of soy and sunflower mixed flours may complement each other [ 70 ]. According to Osen et al. [ 32 ], extrusion did not influence hydrolysis or amino acid composition, possibly because high feed moisture minimized shear stress and mechanical energy loss in the extruder. The main amino acids in the WG/SPI meat analogues were glutamic acid, proline, leucine, and aspartic acid, according to Chiang’s study [ 22 ], and the amount of cysteine in the meat replacements was higher than that of firm tofu and steamed chicken. It has also been discovered that adding green tea to TVP improved texturization and antioxidant properties but had a negative effect on the expansion and NSI [ 90 ]. Recently, Sakai et al. [ 26 ] and Chen et al. [ 21 ] investigated the in vitro gastrointestinal digestibility of meat analogues. It was reported that HME could improve protein digestibility in several protein materials [ 21 ]. It seems that the extrusion field in this phase focused on finding approaches to mimic the structure of the meat. The next phase will also consider the amino acid profile or protein quality (PDCAAS), but we are not there yet. This follows a logical order, since there is no need to optimize the nutrition profile if the materials do not fulfill the texture criteria.

Cooking Quality

Cooking quality in this review was related to analyses performed before and after extrusion, such as cooking loss or frying loss, swelling index, water absorption capacity (WAC), and breakage rate. The percentage difference between the weight of the sample before and after cooking is referred to as “cooking loss”, and it is an essential indicator in determining the quality of the meat analogues in relation to the amount of juice it retains and the amount of product it produces overall [ 26 ]. In general, preparation factors such as composite materials affect cooking loss in processed meat products [ 50 ]. For instance, in mushroom sausage, the addition of all types of binding agents decreased cooking losses, with carrageenan giving the best results, followed by xanthan gum, soy protein concentrates, and casein [ 66 ]. According to Neumann [ 95 ], the water absorption capacity of a product correlated with its texture after rehydration. Lin and colleagues [ 62 ] studied water absorption capacity in the extruded meat analogues and discovered that samples extruded at the high moisture content (70%) and high cooking temperatures (149 °C and 160 °C) had the highest WAC. Furthermore, the authors reported that the porosity of meat analogues influenced the WAC, since extrudates with comparable physical structures and similar moisture contents did not significantly vary in WAC.

Sensory Evaluation

Another important test that many researchers employed to confirm that the products are acceptable from the consumer’s perspective is the sensory evaluation. The sensory qualities of the generated meat analogues can be evaluated using descriptive sensory analysis and the hedonic scale. A scale or a test using hedonic 7-point, 9-point, and 11-point scales was used to achieve this by untrained, semi-trained, or trained panelists. De Angelis et al. [ 49 ] used an 11-point structured scale ranging from 0 to 10, and the sensory assessment emphasized a powerful odor and taste profile of dry-fractionated pea protein and oat protein (PDF–OP), whereas the extrudates generated by protein isolates had neutral sensory features. It was found in this review that sensory assessment is not included in many investigations of meat analogues; most of them are studies on restructured meat substitutes. Sensory evaluation was conducted by Grahl et al. [ 76 ] using conventional profiling on spirulina and soy-based meat analogues to measure the intensities and amplitudes of the feelings as well as to subjectively characterize the samples. Rousta et al. [ 56 ] evaluated the texture of patties manufactured from Aspergillus oryzae biomass (edible fungi) and compared them with two other commercial patties in Sweden, namely, Beyond and Quorn. The study also revealed that restructured meat could give a different taste because of the chemicals and enzymes used in the pre-treatments, which degrade carbohydrates, proteins, and fats into parts [ 56 ].

Other Assessments

In this sub-theme, several assessments were found to be used in the reviewed studies, such as volatile compounds [ 52 ], microbiological evaluation of meat analogues [ 75 ], and life cycle assessment of products produced from meat analogues [ 104 ]. Kaleda et al. [ 52 ] found that extrusion decreased volatile compounds due to high temperature (150 °C). The microbiological evaluation of meat analogues products was discovered to be a crucial assessment to regulate the microbial growth of the meat analogues and at all stages of processing as well as to estimate the product’s shelf life. Filho et al. [ 75 ] analyzed the properties of raw materials, evaluated the microbial limit testing of the canned product before retorting it, and investigated the most critical processing stages to exert control over the growth of microorganisms. However, to go on to the next phase, researchers must first complete the assessments described above, and that will provide a good understanding of each material and the fundamental processing approach.

4. Recommendation for Future Studies

The findings of this review made it evident that a significant amount of research has been conducted on developing better meat substitutes by using improved texturization methods. The recent trend in publishing indicates that technology in developing meat analogues is receiving greater attention. Even though methods involving extrusion are being employed on an industrial scale, these processes are still in their formative stages and have great room for improvement. Fiber and protein–protein interaction research were less prevalent than in other subjects. However, several attempts were found, such as morphology development (cryo-imaging) and flow characteristics (closed-cavity rheometer) with online sensors and simulations showing a promising future in this area. For high-moisture extrusion technology to be used efficiently, researchers need to have a solid understanding of how changes in the conformation of macromolecules occur and an understanding of the mechanism behind it. This is due to a limited number of studies that have been conducted to evaluate and understand the texturization methods that are now in use. Regarding textural features, researchers should consider what texture they should aim for, including the cooking methods. It is also necessary to conduct more studies on physical structuring approaches that have shown potential, such as extensional shearing devices and high-pressure processing, to diversify methods or improve existing methods.

With the passage of time and the growth of knowledge, several new start-up businesses such as Beyond Meat, Meatless, and Impossible Foods, as well as some popular brands such as Ikea, are concentrating on capturing the meat analogues market worldwide. In this review, sensory assessment and nutritional research on meat analogues were less common than in other sub-themes. Sensory experiments on novel plant-based protein meat analogues might be carried out to determine the market acceptance of each product. Despite the variety of novel plant proteins, additional knowledge is required on the protein powder components that govern extrudability. Future studies on the absorption rate in the gastrointestinal system should also be conducted to ensure that the human body can absorb the nutrients in the produced meat analogues.

5. Conclusions

In this review, we conducted a systematic review and bibliometric analysis of the literature on 84 articles published between 1972 and January 2022. Two research questions were addressed: (RQ1) To what extent has research been conducted on the development of meat analogues derived from plant protein-based materials? The response to this question was presented as a set of findings in Section 3.1 , Section 3.2 and Section 3.3 in descriptive analysis; (RQ2) What are the key research themes in the literature on plant-based meat analogues? The answer to this question was given in the discussion part of Section 3.4 , where we divided our findings into six different themes: (i) objectives of the study; (ii) type of plant protein; (iii) product type; (iii) added ingredients; (iv) texturization technique; and (v) quality assessment.

While this review study has been conducted comprehensively, there are several practical limitations, as mentioned in other studies. Despite the fact that Scopus and WoS are two of the most widely used databases, there are still many journals that have not been indexed. Even though we employed a broad search string, it is possible that some studies were missed by our review. There is a chance that not all journals were included in the search, since no search term is 100% accurate. In future literature reviews, using additional search engines or databases may benefit the broad overview. In addition, this study did not include languages other than English, which might be seen as a source of bias, particularly for publications conducted in non-western cultures.

This study shows how research on meat analogues shifted from focusing on primary components, principally soy, to novel protein alternatives, complete and partial, and lastly to more advanced materials. It was apparent how research moved from individual relationships between protein to multidimensional and integrative research on protein and its chemical changes and structures, and protein–protein interactions during thermal–mechanical processing. This advancement permitted the inclusion of a more extensive range of issues based on plant protein beyond animal flesh. Future studies should focus more on fibers, protein–protein interactions, and macromolecule conformations and mechanisms, diversifying or improving current methods, sensory attributes, and the gastrointestinal absorption rate of each novel protein ingredient.

Acknowledgments

The authors thank Noor Faizawati Badarudin from the Department of Design Sciences, Lund University, for advice and assistance in providing examples of information.

Author Contributions

Conceptualization, I.Z., K.Ö., J.K.P. and M.R.; methodology, I.Z., K.Ö., J.K.P. and M.R.; validation, I.Z., K.Ö., J.K.P. and M.R.; formal analysis, I.Z.; investigation, I.Z.; data curation, I.Z.; writing—original draft preparation, I.Z.; writing—review and editing, I.Z, K.Ö., J.K.P. and M.R.; visualization, I.Z.; supervision, K.Ö., J.K.P. and M.R. All authors have read and agreed to the published version of the manuscript.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Conflicts of interest.

The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

Funding Statement

This research was funded by the Malaysian Agricultural Research and Development Institute (MARDI), grant number 20190815.

Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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• Published: 07 August 2024

Advancements in plant based meat analogs enhancing sensory and nutritional attributes

• Jiwon Jang 1 &
• Dong-Woo Lee   ORCID: orcid.org/0000-0002-2272-8321 1 , 2  

npj Science of Food volume  8 , Article number:  50 ( 2024 ) Cite this article

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• Biomaterials – proteins

The burgeoning demand for plant-based meat analogs (PBMAs) stems from environmental, health, and ethical concerns, yet replicating the sensory attributes of animal meat remains challenging. This comprehensive review explores recent innovations in PBMA ingredients and methodologies, emphasizing advancements in texture, flavor, and nutritional profiles. It chronicles the transition from soy-based first-generation products to more diversified second- and third-generation PBMAs, showcasing the utilization of various plant proteins and advanced processing techniques to enrich sensory experiences. The review underscores the crucial role of proteins, polysaccharides, and fats in mimicking meat’s texture and flavor and emphasizes research on new plant-based sources to improve product quality. Addressing challenges like production costs, taste, texture, and nutritional adequacy is vital for enhancing consumer acceptance and fostering a more sustainable food system.

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Introduction.

Over recent decades, the global meat industry has experienced substantial growth. By 2020, meat production reached around 337 million tons, nearly five times the amount produced in the 1960s 1 . Europe and North America initially led production, but by 2020, Asia became the major contributor, accounting for 41% of global meat production (Fig. 1a ). This growth is attributed to the doubling of the global population and significant socio-economic changes. Projections suggest that by 2050, as the global population approaches 9 billion, demand for meat production could increase by 50–73% 2 . This trend is due to population growth and a tripling in global income over the past fifty years, making meat more accessible and increasing its consumption 3 (Fig. 1b ). The livestock sector, including meat and dairy, significantly impacts the environment, contributing to deforestation, greenhouse gas (GHG) emissions, and water pollution 4 . Additionally, high red meat consumption is linked to health risks like cardiovascular diseases, colorectal cancers, and type 2 diabetes 5 . Conventional meat production also raises animal welfare concerns, often involving inhumane practices 6 .

a Global mean meat production from 1960 to 2020. In 1961, 12 (7.3%) of 164 countries produced more than a million tonnes of meat, whereas by 2020, 46 (10.9%) of 193 countries surpassed this production milestone. b The global average meat consumption per capita from 1960 to 2020. In 1961, 5 (2.0%) of 250 countries had a meat consumption of at least 100 kg/year per capita, accounting for 0.037 billion people. By 1990, 9 (3.42%) of 263 countries reached this consumption level, corresponding to 0.32 billion people. In 2020, 9 (3.57%) out of 252 countries reported per capita intakes of at least 100 kg, representing 1.06 billion people. Source: Our World in Data ( https://ourworldindata.org ).

The meat consumption and production challenges are multifaceted, encompassing food security, health risks, and animal welfare concerns 7 . About 800 million people globally face chronic hunger, with another 2 billion suffering from micronutrient deficiencies 8 . The COVID-19 pandemic highlighted vulnerabilities in the food supply chain, underscoring the need for resilient and sustainable food systems 9 . While meat is a vital nutrient source, excessive consumption, mainly processed meats, is associated with various health complications 10 . Acknowledging these unsustainable practices, the Food and Agriculture Organization (FAO) advocates a shift towards plant-based diets to mitigate climate change impacts and promote sustainable food ecosystems 11 . Innovative alternatives like plant-based substitutes and cultured meats have emerged, offering more sustainable and ethical food choices. Plant-based meat analogs (PBMAs) have gained attention for their potential to provide a more nutritious profile than red meat. These alternatives aim to replicate the texture and flavor of animal meats using plant-derived ingredients, but they face challenges in authentically replicating conventional meat 12 . This review seeks to navigate these challenges, examining the latest trends, ingredients, and innovative methods in the meat alternative sector.

Types of alternative meats

Alternative meats encompass four primary categories based on their origin: plant-based, microorganism-based, animal-based, and insect-based meat analogs, each with distinctive sources and production processes (Fig. 2 ).

This infographic delineates the production process of different types of meat alternatives.

Plant-based meat analogs (PBMAs)

PBMAs, crafted mainly from legumes, soybeans, wheat, and lentils, mimic the fibrous texture of meat using protein reforming techniques like extrusion, shear cells, and three-dimensional (3D) printing 13 . These analogs, considered healthier and more eco-friendly due to lower calorie and saturated fat content, still face challenges in preserving nutritional value and cost-effectiveness during intensive processing, which may lead to increased salt content and reduced essential micronutrients 14 . Nevertheless, PBMAs are gaining commercial momentum, with ongoing production methods and raw material selection improvements to enhance their sensory and nutritional appeal.

Microorganism-based meat analogs (MBMAs)

MBMAs, particularly mycoprotein from fungi like Fusarium venenatum , offer rich protein and fiber content and notable health benefits 15 . Production involves using agricultural and industrial by-products and crucial freezing stages for meat-like texture 16 . Despite challenges like resource-intensive production and allergenicity, microalgae like Chlorella and Spirulina are gaining attraction in PBMA applications due to their rich nutrient profiles and functional properties 17 . However, they face hurdles related to color, odor, and texture in PBMA integration 18 .

Animal cell-based meat analogs (ABMAs)

Lab-grown meat, an eco-friendly alternative, is produced by cultivating stem cells from muscle tissue or embryos in bioreactors 19 , structured on edible scaffolds 20 , and formed into muscle fibers using tissue engineering techniques 21 . Since the first lab-grown hamburger debuted in 2013, significant advancements have been made in scaffold technologies, including 3D bioprinting 22 and electrospinning 23 . Despite its potential to mimic meat’s sensory attributes and recent regulatory milestones 24 , cultured meat still faces challenges in fat composition modification 25 , nutrient content adjustment 26 , technical complexities like co-culturing different cell types 27 , scalability, production costs, and public acceptance 28 .

Insect-based meat analogs (IBMAs)

Insect proteins from crickets, mealworms, ants, and black soldier flies offer a promising alternative rich in nutrients like amino acids and bioactive peptides 29 . Despite their potential and global inclusion in diets, concerns about safety, shelf-life, and consumer acceptance necessitate stringent standards, thorough quality checks, and proper regulation. Addressing health risks, ensuring safe preparation 30 , and devising strategies to increase consumer acceptance, such as incorporating insects into familiar foods or inconspicuous powdered forms, are pivotal for advancing IBMAs 31 .

Classification and evolution of PBMAs

The development of PBMAs has seen significant advancements over time, evolving through several generations of products and innovations (Fig. 3 ). Initially dominated by soy protein, the sector expanded to diverse plant-based proteins, leveraging ingredients such as pea, wheat, potato, mung bean, and rice (Table 1 ). This journey from traditional products to advanced alternatives highlights the plant-based meat industry’s commitment to addressing consumer preferences and environmental concerns, continually evolving to overcome earlier limitations.

This infographic provides a detailed view of the PBMA production steps. The left side details the key ingredients: plant-derived proteins that shape texture and flavor, flavoring agents that enhancing taste while masking off-flavors, and texturizing agents for meat-like fibrous structures. The right-side outlines manufacturing phases, from upstream protein optimization and various structuring techniques to create meat-like textures, to final formulation steps integrating binders, flavors, colorants, and nutrients.

The 1st generation of plant-based meat

Initially, the PBMA sector relied on traditional Asian protein sources like tofu and tempeh, known for their unique textures and rich nutrient profiles 32 . Tempeh from Indonesia, for example, effectively emulates meat texture, while yuba, a Japanese culinary product, serves as an eco-friendly wrapping material 33 . Innovations such as textured vegetable protein (TVP), seitan, and mycoprotein emerged to meet the rising demand from vegetarians and those preferring plant-based diets. TVP, processed via low moisture extrusion (LME), is designed to achieve a meat-like texture when rehydrated 34 . Seitan, made from vital wheat gluten, offers a chewy texture similar to meat. Mycoprotein, marketed as Quorn by Marlow Foods in 1985, quickly gained popularity as a meat substitute 35 . These early products, foundational to today’s robust plant-based meat industry, often lacked the complex texture and flavor diversity of animal meat, offering a somewhat limited sensory experience 34 . For example, replacing 30% (w/w) of beef with TVP in sausages adjusted the water content, brightness, and yellowness, decreased fat content and hardness, thereby affecting sensory attributes 36 . Additionally, the texture of PBMA patties, characterized by lesser hardness, chewiness, and gumminess compared to real meat, was to some extent improved by incorporating additives like methylcellulose to boost taste and flavor 37 .

The 2nd generation of plant-based meat

The second generation of plant-based meats aimed to more closely replicate the fibrous texture and flavor of animal-based meats, leveraging advanced technologies such as high-moisture extrusion (HME), shear cells, and 3D printing 38 . These approaches range from bottom-up methods such as microbial fermentation 39 , electrospinning 23 , and cultured meat 22 , which integrate aligned fibers with various ingredients, to top-down methods including extrusion 40 , freezing 41 , and shear technology that create fibrous textures by stretching biopolymer blends 42 . Bottom-up methods often face challenges in accurately replicating muscle structure and achieving scalability. Conversely, HME has proven highly effective, employing intense mechanical mixing under high pressures and temperatures to transform plant-based protein isolates into products that closely mimic the texture and flavor of meat 43 . The HME process involves several stages within the extruder (Fig. 4a ): starting with a low-temperature mixing zone that hydrates proteins without altering their structure, moving to a moderate-temperature zone for partial denaturation, and culminating in a high-temperature melting zone where proteins fully denature 44 . Controlled cooling solidifies the melt, facilitating phase separation and bond formation to finalize the fibrous structure 45 . Leading brands like Marlow Foods, Gardein, Beyond Meat, and Impossible Foods have continuously refined their offerings, significantly narrowing the gap between plant-based and traditional meat products.

a Structuring technologies are employed to emulate meat-like texture and shape. b Schematic representation of the manufacturing process for PBMAs using extrusion.

The 3rd generation of plant-based meat

The third generation emerged in response to environmental concerns and a surge in veganism, particularly during the COVID-19 pandemic. This generation debunks myths about plant-based meats being inherently less healthy and emphasizes high-quality, minimally processed products with clear labeling and fewer additives. Despite facing market fluctuations and competition challenges, innovation and adaptability to shifting consumer needs remain crucial for the ongoing success of the plant-based protein sector 46 .

Characteristics of plant-based meat ingredients

PBMAs harness biochemical similarities between plants and animals, with primary raw materials comprising proteins (20–50%), polysaccharides (2–30%), and fats (0–5%). These components are critical in determining the texture, while additional ingredients enhance flavor, color, and nutritional value. Effective PBMA formulation is paramount for obtaining desired texturization, binding, and dietary outcomes. In typical PBMA formulations, a blend of proteins, polysaccharides, and fats mimics the texture of meat 47 . Core ingredients include isolates and concentrates from soy, peas, and wheat. Recent developments have introduced alternative proteins such as potatoes, rice, lentils, and algae, improving texture, flavor, and nutrition 14 . Moreover, combinations like soy or pea proteins with cereal proteins (e.g., brown rice) improve the amino acid profiles 48 and promote varied structural properties 49 , enhancing the overall quality of PBMAs.

Proteins in PBMAs

Plant-based proteins significantly influence meat analogs’ structure, color, texture, and flavor, going beyond their nutritional role due to technical properties like solubility, emulsification, foam formation, and gelation 18 . Soy and pea proteins are prominent due to their cost-effectiveness, availability, and processing flexibility. They are often blended with wheat, potato, mung bean, and rice proteins to enhance nutrition and texture (Table 1 ). These plant-derived proteins, categorized into albumin (water-soluble), globulins (salt-soluble), prolamins (alcohol-soluble), and glutelins (soluble in weak acid/alkali solution), contribute to functional and nutritional qualities 17 .

Soy protein, a top choice in PBMAs, effectively mimics meat texture but may face issues like aftertaste and allergenicity. Derived from soybeans containing around 40% crude protein 50 , soy protein powder is processed through cleaning, grinding, dehulling, and de-oiling steps. Soy protein concentrate (SPC) and isolate (SPI) result from extracting soluble and insoluble carbohydrates from defatted soybean flakes, offering varying protein content levels 51 .

Pea protein is gaining prominence in the PBMA protein domain for its processing attributes 52 and mild taste, rich protein quality, and amino acid profile 53 . However, it faces challenges in gel formation compared to soy protein. This disparity presents obstacles during the texturing phase of PBMA production, a crucial step in achieving the desired consistency and feel of the final product 54 . Innovative solutions like enzymatic modifications and polysaccharide conjugation are being explored to improve its techno-functional properties, such as gelling, emulsification, water retention, and oil retention 55 .

Wheat protein, known as gluten, is favored in PBMAs for its water retention capability, enhancing product juiciness and tenderness. Formulating PBMAs with wheat gluten involves blending it with other plant proteins, improving fibrous texture through protein cross-linking, and enhancing digestibility in extruded products 49 . This combination effectively leverages the nutritional synergy between grains and legumes, providing a comprehensive array of essential amino acids vital for human health 56 .

Potato protein, derived from potato juice, is recognized for its emulsifying properties and low allergenic risk, making it a versatile and appealing option for PBMA formulation, especially for those with dietary sensitivities 57 .

Rice protein, valued for its hypoallergenic traits and balanced amino acid profile, faces challenges in water solubility and gelling capability, which are being addressed through ongoing research to enhance its role as a texture modifier in PBMAs 58 .

Pulses are attracting significant attention as a burgeoning source of protein, offering a wealth of nutrients, including proteins, complex carbohydrates, vitamins, and minerals 59 . Lentil protein, praised for its gelling, emulsifying, and foaming properties, mirrors the functional traits of pea and soy proteins. Mung bean proteins excel in gel formation and water retention, albeit with variability in amino acid composition 60 . Faba beans stand out for their superior biological nitrogen fixation, promising enhanced agricultural productivity, reduced environmental footprint, and improved cost-efficiency, thus positioning them as a substantial addition to plant-based protein sources 61 .

Oilseed proteins, a by-product of edible oil processing, are emerging as eco-friendly options in the food industry due to their minimal anti-nutritional factors, abundance of essential amino acids, and high digestibility 62 . Notably, chia, hemp, and pumpkin seeds have been recognized for their health-promoting properties, including anti-inflammatory and cardioprotective effects, making them popular in health-focused food formulations 63 . An example of their applications is found in products like the Whole Burger from the plant-based meat brand Abbot’s, which incorporates chia seeds ( https://vegoutmag.com/ ). This burger is reported to contain 22 g of protein per serving, potentially offering greater satiety and muscle health support compared to traditional veggie patties.

Pseudocereals like amaranth and quinoa are esteemed for their high-quality proteins loaded with essential amino acids and excellent gelling properties. These proteins significantly enhance the nutritional value and texture of PBMAs. Amaranth is renowned for its broad health benefits, while quinoa seeds are valued for their abundance of health-promoting flavonoids and antioxidants 64 . Isolated quinoa protein, in particular, is acclaimed for its superior emulsifying and gelling capabilities, broadening its application in food product development 65 .

Algae proteins, rich in essential amino acids and beneficial omega-3 fatty acids, are valued for their nutritional benefits in food products 66 . A New York-based alternative meat brand, Akua, utilizes sustainably farmed kelp to produce the Kelp Burger, an innovative addition to their product line which includes items like kelp jerky, known for its crispy exterior and warm interior ( https://settingmind.com/ ). However, while seaweed-derived polysaccharides enhance texture, their unique sensory attributes can limit their application in specific food applications 67 .

Aquatic plants like duckweed ( Lemna minor ) are notable for their substantial protein content and rapid biomass production, closely mirroring the nutritional profile of animal proteins and often surpassing traditional plant proteins 68 . Duckweed is especially valued for its ribulose 1,5-bisphosphate carboxylase (RuBisCo) content, a crucial component known for its desirable properties and role as a precursor to bioactive peptides. However, the challenge lies in optimizing the extraction and isolation techniques for RuBisCo to fully leverage duckweed’s potential in food applications 69 . Meanwhile, products like Rubi Whisk™ have already harnessed similar plant-based advantages. Marketed as an egg replacer for PBMAs, Rubi Whisk™ offers enhanced fat binding, oil holding, and water-binding capabilities, providing a clean label, plant-based solution that enhances the structural integrity, moisture, and oil retention of allergen-free baked goods ( https://www.plantiblefoods.com ).

Polysaccharides in PBMAs

Polysaccharides play a pivotal role in enhancing the functional and structural aspects of PBMAs. Derived from sources like potato, maize, wheat, cassava, pea, and rice, polysaccharides, including starches and flours, act as efficient fillers, improving texture and ensuring consistency across PBMAs. Fibers from peas, potatoes, and bamboo, along with polysaccharide gums such as xanthan gum and carrageenan, are instrumental in thickening PBMA products and reducing cooking loss. Their ability to retain water excellently and to form stable oil/water emulsions is crucial for attaining the desired consistency and mouthfeel in PBMAs 70 .

Incorporating dietary fiber into PBMAs enhances their nutritional value and imparts critical functional attributes. The capacity of dietary fiber to hold water is vital in mimicking the structure and texture of meat 71 . Including dietary fiber in PBMAs can lead to an increased fiber intake, potentially yielding health benefits like lowering LDL cholesterol and reducing risks associated with cardiovascular disease and obesity 72 , 73 .

Starch is a fundamental functional component in PBMAs, significantly impacting product yield, moisture retention, and texture modification 74 . Starch varieties with high amylopectin content, such as wheat and maize starch, are particularly effective in lending softness to PBMAs during the extrusion process, a vital characteristic for replicating the tenderness and bite of meat 75 . As such, starch is integral to PBMA formulations, serving as a primary structuring agent and significantly contributing to the texture and overall quality of the final product.

Fats in PBMAs

Fats are essential in PBMAs for replicating traditional meats’ juiciness, tenderness, and flavor profiles. During heat processing, the onset of lipid oxidation plays a critical role in developing flavor compounds that enhance the meat-like taste 76 . In PBMAs, various plant-based oils, including coconut, sunflower, and avocado oils, are frequently used due to their health benefits compared to animal fats 70 . The desired characteristics of the final product guide the selection of these oils. Balancing unsaturated and saturated fatty acids is crucial in replicating the sensory qualities of meat. For example, semi-solid oils such as coconut oil can replicate the appearance of meat marbling. Still, their higher saturated fat content requires careful consideration to maintain the healthfulness of PBMAs 72 .

Techniques for crafting PBMAs

The production of PBMAs involves a sophisticated process of protein extraction from sources like soy, pea, and wheat, structured into meat-like textures using techniques such as extrusion, shear cells, spinning, and 3D printing (Fig. 4a ). Flavoring agents, fats, and additives are introduced to enhance the sensory appeal, with binding agents, texturizers, flavors, and coloring agents playing crucial roles in product enhancement.

Structuring techniques

Plant-derived proteins are processed into forms that mimic the texture of traditional meat through various manufacturing techniques, notably LME and HME. LME transforms raw materials into a semi-solid blend, which is thermally treated in a cooking zone and then extruded through a long, slender die. This process produces TVP, characterized by an extended shelf life and a meat-like fibrous texture upon rehydration (Fig. 4b ). LME typically results in greater extrudate expansion compared to HME. On the other hand, HME requires higher moisture content (40–70%) and an extended cooling die, creating layered, fibrous textures resembling whole-cut meat 40 . During HME, proteins are exposed to high temperatures, shear forces, and pressures, which facilitate the formation of cross-links and new protein-protein associations, ultimately yielding layered, fibrous textures that closely resemble the structure of muscle meat 77 . Additionally, optimal HME processing requires careful adjustment of various parameters, including the rotating speed of screw, barrel temperature, and feeding amount to achieve the desired textural outcomes. HME products, typically maintained in refrigerated or frozen states, closely resemble muscle meat in texture and appearance 78 .

Advanced structuring techniques such as 3D printing and shear cell technology enhance sensory experiences and nutritional value. Shear cell technology, still in its pilot phase, creates unique product structures by inducing flow within the product, resulting in distinctive fibrous textures. It encompasses phases similar to HME, including amalgamation and hydration, thermo-mechanical treatment, and cooling. While both HME and shear cell technology utilize shear forces to deform and align biopolymers, the resulting structures differ significantly in macroscopic appearance 42 . Extruded meat analogs exhibit a V-shaped pattern, where individual layers align parallel to the die wall. In contrast, the macrostructure of products obtained through shear cell technology displays distinct fibers oriented along the direction of the applied shear flow. Also, shear cell technology can potentially create PBMAs with greater thickness than those produced by HME methods 79 .

3D printing constructs products layer by layer, thereby enhancing sensory experiences and nutritional value 80 . This technology utilizes bio-inks derived from various animal and plant cells to fabricate cultivated meat (CM) that authentically mimics meat textures. It combines plant-based proteins and fibers using methods such as extrusion 3D printing, inkjet printing, and binder spraying, which are essential for developing plant protein-based edible inks from sources like soy protein isolate and wheat gluten 81 , 82 . For instance, an air-heating extrusion-based 3D printer uses a balanced blend of soy, wheat, and rice protein pastes to improve printing performance and create a meat-like layered structure 83 . Despite its potential, 3D printing faces challenges such as imprecision and suboptimal productivity, necessitating intensive research for successful commercialization 80 . Technical issues such as low viscosity, ink stability, nozzle clogging, and achieving adequate mechanical integrity are being addressed through technological improvements in extrusion techniques and material formulations. These enhancements aim to enhance the physicochemical properties of the final products 84 . Startups like Revo Foods and Nova Meat are utilizing extrusion 3D printing and integrating tissue engineering to hasten the development of both plant and cell-based meats 85 . Innovations in coaxial and dual extrusion are being developed to produce multi-material products that resemble whole-cut steaks, although challenges with ink flow characteristics remain 86 . Ongoing research efforts focus on optimizing formulations and nozzle sizes during 3D printing and cooking to enhance product attributes such as hardness, springiness, cohesiveness, and chewiness 87 , 88 . Additionally, blending technologies based on protein-protein interactions have been developed to improve fiber formation, thereby enhancing the rheological properties of bio-inks and boosting their performance in 3D printing applications 83 .

Additionally, the integration of scaffold materials such as edible microcarriers and fiber structures, which are often created using advanced manufacturing techniques, is crucial for tissue development and effective nutrient and oxygen transport within thicker structures 89 . To improve scaffold functionality, modifications like crosslinking with food-grade agents 90 and the incorporation of bioactive polymers such as silk fibroin and gellan gum are used to enhance mechanical properties and cell adherence 91 . TVPs, derived from soy by-products, are used as scaffolding materials due to their structural versatility and nutritional benefits 20 , 92 , supporting efficient cell seeding and differentiation necessary for large-scale CM production.

Innovations in scaffold materials focus on reducing material costs and capital expenses while ensuring safety and optimizing bioprocessing systems for market readiness. As CM progresses towards commercial viability, integrating scaffold materials that minimize environmental impacts and align with consumer sustainability and ethical preferences is increasingly important 93 . Future directions will likely concentrate on enhancing scaffold interactions with cultured cells, reducing reliance on animal-derived materials, and aligning with sustainable food production goals. These initiatives aim to make CM more economically viable and environmentally sustainable, reflecting growing consumer demands for ethically produced alternatives.

Scaffold-free technologies like cell sheet engineering, utilize biocompatible materials (e.g., chitosan, alginate, gelatin) to enhance structural and mechanical properties and reduce costs by simplifying the cell expansion and harvesting process 94 . Cell sheet engineering allow cells to form monolayers and secrete their extracellular matrix, facilitating the noninvasive detachment and stacking of cell sheets to produce thick, high-density tissues without the need for animal-derived materials 95 . These technologies, combined with 3D printing, enable the integration of microcarriers into 3D printed hydrogels 96 or directly within bioinks 97 . Combining cell sheet engineering with nutrient delivery platforms that incorporate algae-derived proteins and growth factors supports cell growth, reduces medium costs, and minimizes the use of animal-derived supplements 95 .

Manufacturing & quality improvement technique

In PBMA production, the sophisticated amalgamation of plant proteins, various food additives, and state-of-the-art technologies is crucial for enhancing flavor and appearance, with ingredient synergy playing a pivotal role in overall quality enhancement and addressing product deficiencies. Techniques like extrusion transform these ingredients through denaturation and gelatinization, followed by reassembly into a structured network via bond modification such as noncovalent and disulfide bonds. This process typically promotes phase separation, forming molecular aggregates within a continuous protein matrix, essential for mimicking meat-like textures.

Binders are essential for manufacturing intricate meat analogs, enhancing texture, color, flavor, processing quality, and nutritional values. Ingredients such as soy or pea proteins alone may be inadequate for creating viscoelastic networks in emulsion-type meat analogs, necessitating the use of binders to foster cohesive and adhesive interactions, such as hydrogen bonding or electrostatic interactions 98 . These interactions help maintain component cohesion, fortify emulsion stability, curb oil leakage, and adhere to the TVP particles. Common binders like methylcellulose, known for its hydroxyl groups, solidify upon heating and revert to a viscous state upon cooling 99 . Other prevalent binders, including pectin, carrageenan, guar gum, cellulose, xanthan, and locust bean gum, contribute unique rheological properties that influence gelation, thickening, and texture 100 . Edible gum, notably carrageenan, and pectin, are often incorporated into soy protein meat substitutes, enhancing taste, texture, hardness, and chewiness, or electrostatically interacting with soy protein to bestow viscoelasticity and maintain stable physical properties under high-temperature shear 101 .

Achieving palatability is fundamental to PBMA’s market success, yet replicating the complex flavor profile of traditional meat poses significant challenges 100 . Key components such as iron, lactate, and inosine 5’-monophosphate are crucial for mimicking the authentic flavors of raw meat, while cooking-induced Maillard reactions and lipid oxidation further enrich the flavor 102 . PBMAs often require larger amounts of flavor enhancers such as savory yeast extract (YE), nucleic acids, and sugars compared to conventional meat. Spices and herbs are also used to emulate the rich flavors of processed meats and to mask any undesirable tastes from legume proteins 14 . Overcoming the beany flavor associated with plant proteins involves inactivating lipoxygenase and neutralizing flavors through fermentation or β-cyclodextrin 103 . Ingredients such as hydrolyzed vegetable protein (HVP) and YE are crucial in PBMA formulations 104 , with HVP producing a strong meat-like flavor when heated with sugars and yeast autolysis, and YE enhancing umami and kokumi flavors upon heating 105 .

The visual allure of meat analogs is paramount for customer acceptance. While traditional meat’s color is primarily attributed to myoglobin 106 , PBMAs often employ soy protein or gluten, resulting in hues that differ from meat’s expected red or brown. To mimic natural meat colors and marbling, innovations in plant-based pigments and ingredients like beetroot 107 and leghemoglobin 108 are used, although concerns about genetically modified origins are driving the development of safer, cost-effective alternatives to traditional coloring agents like titanium dioxide.

The manufacturing and quality improvement of PBMAs involves integrating various ingredients and technologies to enhance structural, flavor, and visual qualities. This includes optimizing key extruder operating parameters such as moisture content, barrel temperature, and screw speeds, which are crucial for proper texturization 87 . These parameters are part of a complex multi-input-output system 109 that significantly impacts the texture and physicochemical properties of the final products 87 , 110 .

Meat substitution, both partial and complete, often use plant-based ingredients, with some commercial products incorporating animal proteins like dairy and eggs 52 , 92 . Brands like Hungry Jack’s® (v2foods™Australia), Impossible™ Burger (Impossible™ Foods, U.S.), the Beyond Burger® (Beyond meat®, U.S.), and Chicken free chicken® (Sunfed®, New Zealand) use a mix of protein sources (soy, pea, wheat) and carbohydrate (cellulose, methylcellulose, starch) to enhance water and lipid interactions, contributing to a meat-like texture 92 . However, the complexity of extrusion conditions—including high shear and temperature—can denature proteins, which complicates texture prediction and functionality. Additionally, factors like storage conditions, preprocessing, and harvesting variations significantly affect these properties, emphasizing the importance of understanding physiochemical interactions during extrusion 111 .

Health consciousness in PBMA production

The rise of PBMA production has been supported by major conglomerates and fast-food chains, aiming to crater to a health-conscious market. However, maintaining a healthy image for PBMA poses challenges. To sustain their perception as a healthier alternative, a majority express willingness to increase PBMA consumption if their nutritional profile aligns with real meat. The perception of PBMA as highly processed food with potential health risks necessitates the development of new technologies and raw materials to address these concerns and shift consumer perceptions. A meticulous analysis of nutritional components and additives in PBMA can pave the way for healthier and safer alternatives.

Nutritional quality in PBMAs

PBMAs strive to emulate the appearance and taste of meat, but nutritional quality can sometimes be compromised. Generally, PBMA products contain fewer calories, less total and saturated fat, and more dietary fiber than their meat counterparts. However, they may exceed sodium recommendations and lack essential micronutrients like iron, zinc, and vitamin B 12 112 . Addressing these deficiencies through fortification can enhance the health profiles of PBMAs, positioning them as healthier alternatives

The surge in demand for plant-based proteins has spurred in-depth research into protein quality and bioavailability, assessed through the dietary indispensable amino acid score (DIAAS) and the protein digestibility-corrected amino acid score (PDCAAS) 113 . DIAAS measures the true ileal digestibility of essential amino acids in foods, while PDCAAS assesses protein quality based on digestibility and amino acid composition. To improve protein quality in PBMAs, integrating diverse plant sources and processing methods can enhance bioavailability by unfolding proteins, reducing antinutrients, and fostering optimal hydrolysis 114 . Although plant-based proteins often lack essential sulfur-containing amino acids 8 , 52 , HME cooking preserves lysine and enhances the bioavailability of other amino acids through thermal unfolding, making proteins more digestible 115 . Despite high moisture content potentially reducing the effectiveness of antinutritional factors (ANFs) deactivation 116 , the thermal and mechanical forces in HME cooking generally improve digestibility and reduce some ANFs, enhancing the nutritional profile of PBMAs 117 .

Antinutrients such as tannins, phenols, saponins, phytates, glucosinolates, and erucic acid can impede nutrient digestion and absorption 118 . Techniques like fermentation, soaking, and various processing methods (e.g., gamma irradiation, germination, heating, genomic technology, sonication, microwave, high-pressure processing, and electric field methodologies) can reduce their concentration and mitigate adverse effects 119 . For instance, extrusion processes enhance the bioavailability of essential amino acids while reducing ANFs, thereby improving the overall nutritional value of PBMAs.

While plant-based diets are typically low in sodium, PBMA products may have elevated sodium levels due to processing, posing health risks such as chronic kidney disease 120 and increasing lithogenic risks among children and infants 121 . Reducing sodium content through natural seasonings and exploring sodium-curbing flavors and processing techniques can offer healthier PBMA options 122 .

Iron, zinc, and vitamin B 12 are challenging to obtain in a meat-free diet, and their bioavailability in meat substitutes can be limited by factors like phytate content 123 . Addressing these micronutrient deficiencies is crucial, especially among specific demographic groups 124 . Vitamin B 12 , absent in plants, is an additional challenge, causing deficiencies, especially among vegetarians, vegans, pregnant women, or females in their reproductive years 125 . Fortifying PBMA products and advocating for nutrient-rich plant foods in diets can mitigate these challenges 126 . There is a critical need for education and guidelines centered on plant-based nutrition and fortification to ensure healthy and sustainable diets.

The effect of PBMAs on human gut microbiome

The shift towards reduced meat consumption has increased interest in PBMAs, particularly among flexitarians concerned about meat’s association with cardiometabolic diseases and gut dysbiosis. Despite this interest, some consumers hesitate to adopt PBMAs, perceiving them as low-quality and highly processed, potentially harmful to gut microbiota. This underscores the necessity for targeted research into the complex relationship between meat intake and gut microbiota composition, which is crucial for human health 127 .

Recent studies demonstrate that PBMA consumption may beneficially impact the gut microbiome by increasing butyrate production and microbial diversity 128 . These effects are often attributed to the increased dietary fiber in PBMAs rather than their intrinsic properties. However, the limited scope of these studies, often with small sample sizes, necessitates further research with robust methodologies to substantiate these health implications comprehensively. For instance, research using the TIM gastrointestinal model showed that plant-based burgers (PB) influenced the lipemic response and gut microbiome differentially than beef 129 . PBs generally contain higher carbohydrate levels due to the addition of binders like potato starch, methylcellulose, and maltodextrin, which contribute to a meat-like texture, while typically having lower protein and fat content. Short-term PB intake significantly alters the gut bacterial colonization, increasing the Firmicutes to Bacteroidetes (F/B) ratio and affecting short-chain fatty acid (SCFA) profiles, indicating that the nutritional content and structure of food influence gut microbiota composition. However, it is unclear whether these phenotypes are ascribed to the types of meat or the abundance and species of components in PBMAs.

Observational and experimental studies have provided inconsistent results regarding the specific changes in gut microbiota composition at different taxonomic levels due to meat consumption 130 . These inconsistencies highlight the complexity of understanding how meat attributes, such as processing and cooking methods, impact gut microbiota 131 . This complexity underscores the need for well-designed randomized controlled trials and systematic reviews that control dietary variables more precisely, aiming to clarify the impacts of meat and PBMAs on gut microbiota and broader health outcomes 127 .

Clean label emphasis in PBMA

PBMA formulations often include proteins, water, flavors, oils or fats, binders, and colorants. However, the presence of numerous additives to compensate for the functional limitations of eco-friendly proteins can lead to extensive ingredient lists, potentially deterring consumers who prioritize natural and transparent ingredients. The trend towards “clean label” products, emphasizing minimal and recognizable ingredients, is growing 132 . Such products resonate with consumers seeking transparency and naturalness in their food choices. Factors influencing consumer perceptions of “naturalness” include the origin of the food, processing methods, E-number labeling, and the presence or absence of chemicals or artificial additives 133 . While chemical additives like methylcellulose (E461) are commonly used in PBMAs as binders for their thickening and emulsifying properties, the demand for clean-label products has spurred the development of chemical-free alternatives such as pea protein and sugar beet pectin 134 . Hydrocolloids from natural polysaccharides like konjac glucomannan, κ-carrageenan, konjac mannan, and xanthan gum are increasingly used for their gelling, thickening, emulsifying, and stabilizing properties, contributing to the desired textural properties of PBMAs 133 . Regulatory changes, such as the European Commission’s ban on certain additives, have prompted the food industry to seek safe, natural whitening ingredients for PBMA formulations. Companies are simplifying ingredient lists and using HME and proprietary processes with multi-functional proteins to enhance the texture and sensory qualities of PBMAs, aligning with the clean label trend and catering to consumer preferences for natural and transparent products. Beyond Meat’s IV platform exemplifies the evolution of PBMA into the third generation. It replaces traditional fats with avocado oil to reduce saturated fat and increase the smoke point, making it suitable for BBQ. Additionally, it cuts sodium by 20% and boosts nutritional value with new ingredients like red lentils and fava bean protein. This innovation has earned it the distinction of being the first plant-based meat to receive clean label project certification.

Challenges and opportunities in PBMA development

Taste is crucial in PBMA purchasing decisions, and enhancing the quality of plant-based proteins can reduce off-notes and improve flavor. However, replicating the texture of animal-based meat poses significant challenges, particularly as some PBMAs lack essential amino acids and micronutrients. Addressing ANFs is vital to enhance the nutritional profile of these products. PBMA production costs currently surpass those of traditional meats, affecting affordability and consumer adoption. Recent advances include the Feature-Augmented Principal Component Analysis to model the extrusion process precisely 135 . This approach has produced six mathematical models and the first visualization software for HME, enhancing equipment setup, parameter adjustments, and quality control.

Moreover, PBMAs provide a sustainable alternative to traditional meat, requiring fewer natural resources and producing less environmental waste. As production scales up and technologies evolve, costs are expected to decrease, making PBMAs more accessible and reducing their environmental impact. Despite growing popularity, continuous improvement in taste, texture, and nutritional value is necessary (Fig. 5 ). Effective marketing and educational campaigns can enhance consumer awareness and acceptance. By tackling challenges like undesirable flavors and nutrient deficiencies, the industry can improve consumer perceptions and contribute significantly to a sustainable food ecosystem.

This infographic presents a step-by-step guide outlining the critical criteria for each phase of PBMA manufacturing and distribution.

Data availability

All data supporting the findings of this study on global meat production and consumption trends are openly available at https://ourworldindata.org .

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Acknowledgements

This work was partly supported by a grant (2021M3A9I4021431), the Bio & Medical Technology Development Program of the National Research Foundation (NRF) of Korea, funded by the Ministry of Science and ICT (MSIT), by a grant (HP23C0158), the Korea Health Technology R&D Project, funded by the Ministry of Health & Welfare, Republic of Korea, and by a grant (200118770), the Technology Innovation Program of Development Program of Korea, funded by the Ministry of Trade, Industry and Energy (MOTIE). Republic of Korea.

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Production of plant-based meat: functionality, limitations and future prospects

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Plant-based meat alternatives have been considered from past years, but recently gained much importance in research development and food industries. Concerns related to health, animal welfare, ethical beliefs and environment act as a driving force for the production of protein-based meat analogues. The major challenging work in the development of plant-based meat is to imitate the texture of meat analogues. The production of plant-based meat analogue needs an astute choice and formulation of constituents to perfectly mimic the fibrous texture of meat. Various plant-based constituents like texturized and non-texturized proteins, fats and oils, thickening agents, binding agents, colouring and flavouring agents used with different processing technologies (extrusion, electrospinning, wet-spinning, shear cell, freeze structuring and 3D-printing) for the production of plant-based meat analogues. This critical review highlights essential ingredients for creating these novel meat analogues, emphasizing protein sources, ingredient functionality, production technologies and health and environmental effects of plant-based meat and consumer acceptance towards plant-based meat.

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Acknowledgements

I would like to express my special thanks of gratitude to my friend Dr. Shahid from South China University of Technology for helping us to finish the wonderful task. After his guidance, I came to know about so many new things about this manuscript.

This work is self-funded. Data avaialability: All data generated and analuzed during this study are included in this published article and its supplementary information files.

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Imran, M., Liyan, Z. Production of plant-based meat: functionality, limitations and future prospects. Eur Food Res Technol 249 , 2189–2213 (2023). https://doi.org/10.1007/s00217-023-04287-w

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Received : 16 February 2023

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DOI : https://doi.org/10.1007/s00217-023-04287-w

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