Multiscale structural effects of ultrasound and pulsed electric field processing on structure–property relationships in tomato systems: a mini-review

1 Introduction

Plant-based foods have highly organised, hierarchical structures that determine their physicochemical, mechanical, and nutritional properties (Joardder et al., 2023). The tomato (Solanum lycopersicum) represents an exceptional model system due to its well-characterised cellular architecture, comprising parenchyma cells embedded in a polysaccharide-rich cell wall matrix that regulates mass transfer and structural integrity (Ordaz-Ortiz et al., 2009). Globally, tomato production exceeds 186 million tonnes annually; over 130 million tonnes are processed, generating eight million tonnes of by-products consisting of skins (61%) and seeds (38%) (Trombino et al., 2021; Mozafari et al., 2024). These residues are valuable secondary raw materials rich in lycopene, phenolics, and dietary fibre, driving circular economy strategies focused on their sustainable utilisation as functional ingredients (Carpentieri et al., 2021; Méndez-Carmona et al., 2022).

Conventional thermal processing alters plant tissues indiscriminately, resulting in undesirable changes to texture, safety, and thermolabile compounds (Jayathunge et al., 2019; Yuan et al., 2026). To address these limitations, non-thermal physical technologies such as ultrasound (US) and pulsed electric fields (PEF) have emerged as alternatives capable of selectively modifying food microstructures. US operates via acoustic cavitation, generating localised mechanical stresses, microjets, and temperature fluctuations that disrupt cell walls and enhance mass transfer (Li et al., 2022). Concurrently, PEF applies short, high-voltage pulses to electroporate cell membranes, increasing permeability and enabling selective transport phenomena without causing significant thermal damage (Toshmamatov et al., 2025).

From a food engineering perspective, the application of these electro-physical and acoustic processes has many more uses than just simple extraction. In the processing of tomatoes, ultrasound (US) and pulsed electric field (PEF) technologies are increasingly being used to improve the efficiency of processes such as peeling, juice extraction and drying, while also enhancing the sustainability of the overall process (El Kantar and Koubaa, 2022). Furthermore, these technologies contribute to food preservation by efficiently inactivating microbes that cause spoilage and degrading enzymes that deteriorate quality, thereby maintaining macroscopic food quality, safety, and structural integrity without the detrimental effects of heat (Toshmamatov et al., 2025).

Concurrently, emerging research has expanded beyond conventional food engineering to examine the biological and physiological responses of living tomato tissues to non-thermal treatments. When applied to living matrices, ultrasound-induced acoustic stimulation and mechanical stress can act as controlled triggers, influencing seed germination and activating physiological stress-response mechanisms (Mozafari et al., 2024; Shafaei Fallah et al., 2025). These physical triggers activate cellular signalling pathways and oxidative stress responses, demonstrating that non-thermal technologies can induce complex biochemical adaptations across multiple biological scales before or during processing. These observations expand the concept of structure–property relationships from processed matrices to living biological systems, where structural perturbations can also regulate biochemical and physiological functions.

Consequently, the tomato serves as a unique multiscale model system bridging green extraction engineering, non-thermal processing, and plant physiology. Within this hierarchical framework, physical stimuli induced by ultrasound (US) and pulsed electric field (PEF) initiate molecular and cellular modifications, including alterations in molecular interactions, membrane permeability, and cell wall organization. These primary events propagate toward the tissue scale by modifying porosity, mechanical cohesion, and mass transfer pathways, which ultimately determine macroscopic properties such as texture, rheological behaviour, processing efficiency, and bioactive recovery. Therefore, this review synthesises recent advances in US and PEF processing of tomato matrices of tomato matrices within a comprehensive multiscale framework, connecting structural modifications from the molecular to the macroscopic level while identifying future research gaps. Literature was selected for its relevance to non-thermal processing, extraction performance, and structural modifications in tomato systems.

As illustrated in Figure 1, US- and PEF-induced physical stimuli initiate specific molecular and cellular modifications, including changes in cell wall organization, membrane permeability, and molecular interactions. These initial alterations propagate toward the tissue level by modifying porosity, structural cohesion, and mass transfer pathways, ultimately determining macroscopic properties such as texture, rheological behaviour, processing performance, and bioactive recovery. Although both technologies enhance functional outcomes, they act through distinct mechanisms: US primarily induces cavitation-mediated mechanical disruption, whereas PEF promotes selective membrane permeabilisation through electroporation.

2 Structural, electrical, and acoustic constraints of the tomato matrix

The response of tomato tissues to non-thermal technologies is strongly influenced by their intrinsic structural and physicochemical characteristics. The tomato fruit is composed predominantly of parenchymatous tissue with a high water content of approximately 93%–95%, thin primary cell walls, abundant intercellular spaces, and a polysaccharide-rich extracellular matrix (Ordaz-Ortiz et al., 2009). These architectural features fundamentally dictate mass transfer phenomena and determine the way external physical forces propagate through the tissue (Joardder et al., 2023). From an ultrasound perspective, the elevated water activity of tomato tissues provides a favorable environment for acoustic wave transmission and cavitation development (Li et al., 2022). The collapse of cavitation bubbles generates localized shear forces and microjets that readily interact with hydrated cell walls and middle lamella structures, promoting tissue disintegration and intracellular compound release (Khalid et al., 2024). However, the presence of internal intercellular air spaces within the parenchyma can act as an acoustic damper, attenuating wave propagation and scattering energy, which requires precise power density adjustments to achieve uniform disruption (Shafaei Fallah et al., 2025). Similarly, the effectiveness of pulsed electric field (PEF) treatments is closely associated with the electrical properties of the tomato matrix. The high moisture content and ionic composition, such as organic acids and mineral salts dissolved in the vacuolar juice, contribute to a relatively high baseline electrical conductivity, enabling efficient distribution of electric fields throughout the product (Toshmamatov et al., 2025). As a result, transmembrane potentials can be generated across cell membranes with relatively low energy inputs, facilitating electroporation and enhanced permeability (Andreou et al., 2020). However, these same conductive characteristics also impose significant engineering limitations. Structural heterogeneities, such as the differences in conductivity between the rigid, insulating outer skin cuticle and the highly conductive internal locular gel, lead to non-uniform electric field distributions (Giancaterino and Jaeger, 2023). Furthermore, high tissue conductivity can cause rapid electrical current spikes, risking an unwanted Joule heating effect that undermines the non-thermal nature of the process (Pataro et al., 2020). Consequently, process optimization requires strict consideration of both the physical principles of the technology and the specific biological architecture of the tomato matrix to ensure homogeneous permeabilization (El Kantar and Koubaa, 2022).

3 Ultrasound effects in tomato systems

US processing relies on the interaction between high-intensity acoustic waves and a liquid medium, generating acoustic cavitation that alters mass transfer and cellular integrity (Li et al., 2022). High-intensity power ultrasound (20–100 kHz) is primarily responsible for process intensification due to its high energy density. The implosive collapse of cavitational microbubbles generates extreme conditions, including high shear forces, microjets (200–700 m/s), and transient temperature and pressure gradients. These phenomena disrupt plant tissue architecture, enhance solvent penetration, and facilitate the release of intracellular compounds, including lycopene, phenolics, and structural pectins (Khalid et al., 2024).

3.1 Multiscale physical and structural mechanisms of US

Recent studies demonstrate that ultrasound-assisted extraction (UAE) operates across multiple hierarchical scales, driven primarily by physical and mechanical disruption:

Molecular Scale: Cavitation-induced mechanical forces facilitate the specific liberation of bound bioactive compounds and antioxidant fractions from the complex tomato matrix, enhancing extract functionality (Li et al., 2022).

Cellular Scale: Cavitation bubble implosion leads to membrane poration and cell wall rupture. This transient permeability enhancement promotes irreversible disruption of cellular compartments in rigid tomato polysaccharide matrices, accelerating intracellular metabolite release.

Tissue Scale: Repeated cavitation causes progressive erosion, particle size reduction, and increased porosity, lowering mass transfer resistance. Marinaccio et al. (2024) confirmed that ultrasound-induced surface erosion and microjetting substantially improve solvent accessibility in tomato skin waste, proving the dominance of physical disruption.

Macroscopic Scale: Cumulative multiscale effects translate into enhanced processing performance. Quantitatively, ultrasound-assisted extraction has been shown to substantially accelerate carotenoid recovery from tomato processing residues, reducing extraction times by up to 50% while maintaining superior extract stability when combined with tailored encapsulation strategies (Li et al., 2022). To systematically monitor and quantify these micro-to-macro structural transitions, food engineers rely on established analytical protocols and profiling tools. The propagation of physical boundaries and cellular structural modifications are validated via analytical platforms, including scanning electron microscopy (SEM) for surface erosion profiling, confocal laser scanning microscopy (CLSM) for membrane integrity assessment, and Fourier-transform infrared spectroscopy (FTIR) to trace chemical alterations in the polysaccharide cell wall matrix (Akhi et al., 2025). These complementary characterization techniques provide valuable evidence of microstructural and chemical modifications induced by ultrasound treatment. However, establishing quantitative structure–property relationships requires integrating these observations with complementary measurements of macroscopic properties, including rheological behavior, texture, permeability, and mass transfer characteristics.

3.2 Biological, biochemical, and physiological responses to US

Following a logical causal cascade, the mechanical stress and localised energy dissipation from US also trigger complex biological responses in living tomato tissues. These responses govern the biochemical profile of the tissues before their physical destruction:

Upstream biological signalling (RNA and enzymes): Acoustic cavitation induces the formation of controlled reactive oxygen species (ROS) and mechanical shearing. In living tomato tissues, these phenomena act as cellular elicitors, upregulating the transcription of stress-responsive genes and activating antioxidant defence enzymes such as superoxide dismutase (SOD), catalase (CAT) and peroxidase (POD) (Mozafari et al., 2024).

Downstream biochemical alterations (metabolites): This enzymatic activation directly modulates metabolic pathways, shifting the synthesis and accumulation of secondary metabolites. Consequently, targeted ultrasound exposure triggers a defensive physiological response that synthesises endogenous protective molecules, significantly increasing the concentration of free phenolics, ascorbic acid, and specific antioxidant fractions within the tomato matrix (Mozafari et al., 2024; Shafaei Fallah et al., 2025).

Macroscopic physiological outcomes: At the tissue and organismal levels, this biochemical shift translates into altered macroscopic properties. For example, ultrasound stimulation modulates metabolic preparation mechanisms in tomato seeds and seedlings, enhancing germination rates and accelerating early seedling growth. It also primes defence pathways, improving overall tolerance to subsequent environmental stresses (Shafaei Fallah et al., 2025).

Therefore, these biological responses should not be considered independent phenomena but rather an extension of the multiscale structure–property framework. Mechanical perturbations induced by ultrasound modify cellular organisation and trigger biochemical signalling pathways, which subsequently influence tissue functionality, metabolic composition, and macroscopic physiological performance.

3.3 Process intensification and parameter sensitivity

UAE performance depends strictly on US processing parameters, including frequency, power density, temperature, time, and solvent properties. Systems operating at 20–40 kHz maximise cavitational intensity, though excessive energy input degrades thermolabile compounds like lycopene. Optimisation studies highlight that moderate temperatures (35 °C–65 °C) and short extraction times maximise efficiency. Li et al. (2022) reported optimised UAE protocols enabling high carotenoid recovery from tomato waste while maintaining compound stability via encapsulation. Solvent selection dictates cavitation dynamics and solute solubility; green solvents like ethanol, ethyl lactate, and natural deep eutectic solvents (NADES) reinforce process sustainability (Badea et al., 2025). Furthermore, Teixeira et al. (2024) demonstrated that UAE systems dramatically enhance the extraction of non-carotenoid bioactives, achieving a 54%–73% increase in anthocyanin recovery from specialized tomato matrices compared to conventional solvent methods.

3.4 Application spectrum in tomato systems

Beyond extraction, US has expanded into waste valorisation, seed priming, and pulp structural modification. In residue valorisation, Solaberrieta et al. (2022) established that UAE significantly improves antioxidant extraction from industrial tomato seed waste, where solvent polarity drives cavitation-induced mass transfer. In seed matrices, ultrasound induces critical microstructural modifications in the seed coat; Memiş et al. (2022) demonstrated that this pre-treatment enhances water uptake and metabolic activation, significantly increasing germination and emergence rates. Similarly, in tomato pulp and puree processing, Hou et al. (2026) proved that ultrasound induces structural loosening, accelerating water migration to reduce drying times, increase matrix porosity, and modify rheological behaviour. Thus, US serves as a multifunctional process intensification technology rather than a simple extraction tool.

3.5 Emerging perspectives and limitations

A major challenge for US scalability lies in yield variability arising from differences in reactor design and processing geometries, which hinders industrial standardisation. Moreover, while cavitation-driven physical mechanisms are well established, their coupling with biochemical responses remains insufficiently understood. Nogueira et al. (2026) revealed that ultrasound exposure triggers complex physiological responses, activating oxidative enzymes and modulating stress tolerance via ROS signalling pathways.

Furthermore, emerging evidence indicating that ultrasound priming can alter drought responses across generations and induce transgenerational plasticity (Farkas and Dobránszki, 2026) shifts UAE from a purely physicochemical engineering tool toward a potential modulator of biological signalling pathways. Future research must integrate multiscale mechanistic models connecting cavitation physics, cellular responses, and macroscopic functional outcomes to enable rational process design within sustainable circular bioeconomies.

4 Pulsed electric field effects in tomato systems

PEF technology applies short high-voltage pulses that induce membrane permeabilisation through electroporation. Unlike conventional thermal treatments, PEF modifies membrane integrity while minimising thermal degradation, making it ideal for plant systems rich in thermolabile molecules (Andreou et al., 2020). In tomato matrices, PEF generates an electric potential difference across cell membranes; when this transmembrane potential exceeds a critical threshold, phospholipid bilayers rearrange, forming pores and increasing permeability (Toshmamatov et al., 2025).

4.1 Multiscale mechanisms governing PEF performance

From a hierarchical perspective, PEF modulates tomato matrices across several dimensions:

Molecular Scale: Electroporation alters intracellular transport pathways and compartmentalisation, directly affecting oxidation kinetics, enzyme-substrate interactions, and bioactive stability.

Cellular Scale: Electro-permeabilisation reduces membrane resistance to solvent penetration, triggering intracellular solute release without destroying the cell wall skeleton.

Tissue Scale: Irreversible electroporation causes progressive structural softening and porosity, loosening the skin-parenchyma boundary to facilitate mechanical peeling and lower mass-transfer resistance.

Macroscopic Scale: Cumulative mechanisms translate into higher extraction yields of lipophilic carotenoids, reduced peeling resistance, lower processing times, and enhanced fresh-like quality retention.

In parallel, the quantification of these electro-physical transitions requires advanced characterization frameworks. Microstructural modifications induced by electroporation are routinely validated via established analytical platforms; specifically, CLSM is utilized to verify cell membrane permeabilization, SEM profiles tissue softening and matrix porosity, and FTIR traces chemical rearrangements within the structural polysaccharide network (Akhi et al., 2025). These analytical approaches provide important insights into membrane permeabilization, tissue organization, and chemical modifications after PEF treatment. Nevertheless, direct quantitative relationships between electrical processing parameters and macroscopic functional properties require additional evaluation through complementary techniques such as texture analysis, rheological measurements, and mass transfer assessments.

4.2 Process intensification and parameter sensitivity

Depending on treatment intensity, electric field strength, and specific energy input, electroporation can be tuned to be reversible or irreversible. Moderate-intensity PEF pre-treatment (0.5–10 kV/cm; 1–10 kJ/kg) effectively permeabilises plant-cell membranes while minimising energy, solvent use, and processing times, facilitating mass transfer, metabolite release, and matrix softening (Belgheisi et al., 2024). Higher field intensities induce irreversible electroporation of microbial cell membranes. Concurrently, electro-physical parameters can induce structural modifications in metabolic enzymes, leading to selective non-thermal inactivation while preserving vulnerable micronutrients like vitamin C (Min et al., 2003). Although PEF is generally considered a non-thermal technology, high electric field intensities (35–40 kV/cm) can generate localized Joule heating within the conductive tomato matrix. Under these conditions, microbial and enzymatic inactivation results from a combined effect of electroporation and localized thermal stresses rather than from purely electro-physical mechanisms.

4.3 Application spectrum in tomato systems

Applications of PEF technology in tomato systems demonstrate how electroporation-driven structural modifications at the cellular level result in changes to tissue functionality and processing performance. Increased permeability of cellular membranes reduces limitations in intracellular mass transfer, facilitating the release of carotenoids and other lipophilic compounds while preserving much of the cell wall architecture. For instance, PEF treatments have been shown to increase lycopene extraction from tomato processing waste by up to 47% (from 9.84 to 14.31 mg/100 g), while also enhancing carotenoid recovery without promoting undesirable cis/trans isomerisation (Andreou et al., 2020; Pataro et al., 2020). Similarly, controlled membrane permeabilisation improved the extractability of lycopene, β-carotene and β-cryptoxanthin by up to 88%, 69% and 24% respectively (Belgheisi et al., 2024).

At the tissue scale, these cellular changes result in reduced mechanical resistance and improved transport properties. PEF-induced softening of the skin–parenchyma interface has been shown to facilitate industrial peeling operations, reducing the required mechanical work by up to 72.3% whilst increasing juice yield by 20.5% (Andreou et al., 2020). Similar reductions in peeling force and improved quality retention have also been reported for PEF-treated whole tomatoes (Giancaterino and Jaeger, 2023). These findings demonstrate that the macroscopic benefits of PEF are direct consequences of electroporation-mediated changes in cellular integrity and tissue organisation.

Beyond fruit tissues, the same mechanism can be exploited in more complex matrices, such as tomato seeds. Here, PEF-induced permeabilisation of lipid-rich structures enhances the extraction of valuable compounds, including linoleic acid-rich oil and associated carotenoids (Toshmamatov et al., 2025). Therefore, the performance of PEF should be interpreted within a multiscale structure–property framework, where controlled membrane modification governs subsequent changes in mass transfer and mechanical properties.

4.4 Emerging perspectives and limitations

Despite combining low thermal impact with targeted structural engineering, PEF presents bottlenecks. Its efficiency depends heavily on matrix electrical conductivity and homogeneity, limiting applications to liquids, purees, or pumpable slurry systems rather than irregular solid masses. Achieving uniform electric field distribution across large-scale chambers remains a challenge, as non-uniformity leads to inconsistent microbial or enzymatic inactivation (Min et al., 2003).

Additionally, transitioning from laboratory batch chambers to continuous industrial co-axial or parallel-plate systems requires complex flow, fouling, and field-control metrics. Future research must focus on coupling predictive electro-fluid dynamics with multiscale structural models to optimise chamber geometries, minimise electrochemical reactions at electrode surfaces, and unlock the commercial potential of PEF within circular agri-food bioeconomies.

To provide a comprehensive comparative overview, Table 1 systematically summarises the multiscale structural effects, operating conditions, and functional outcomes of both US and PEF processing in tomato systems.

TABLE 1. Multiscale structural effects, processing conditions, and functional outcomes of ultrasound (US) and pulsed electric field (PEF) technologies applied to tomato systems.

5 Comparative framework and structure–property relationships

US and PEF processing operate through distinct physical mechanisms but converge toward identical process outcomes. While US induces cavitation-driven disruption of cell walls, middle lamella structures, and tissue architecture, PEF primarily targets phospholipid membranes through electroporation, preserving much of the surrounding cell wall framework (Hou et al., 2026). These architectural changes ultimately govern fluid transport enhancement, mechanical performance, and chemical accessibility within the tomato matrix. Specifically, the disruption of the cell wall and middle lamella induced by ultrasound reduces tissue cohesion, increases porosity and promotes viscosity reduction and rheological changes in tomato pulp. In contrast, pulsed electric field (PEF) processing predominantly modifies membrane permeability while preserving a greater portion of the cell wall architecture. This results in controlled softening, reduced mechanical resistance, and enhanced selective mass transfer. Therefore, the distinct effects of US and PEF on texture, elasticity, permeability, and processing performance are explained by their different structural targets.

Because acoustic processing triggers intense cavitation-induced shear forces, it leads to a more pronounced reduction in viscosity, increased matrix porosity, and altered pulp rheology compared to the selective electro-permeabilisation achieved by PEF (Hou et al., 2026). These distinct microstructural alterations allow food processors to modulate final product texture, thickness, and oxidation susceptibility. In quantitative terms, these structural shifts offer significant processing advantages: optimised UAE protocols can increase carotenoid and lycopene yields from tomato residues by 40%–50% while halving extraction times (Li et al., 2022). Meanwhile, customised PEF applications can boost lycopene extractability by up to 88% (Belgheisi et al., 2024), increase total juice yield by 20.5%, and slash the mechanical work needed for industrial peeling by over 70% (Andreou et al., 2020). To secure the long-term stability of these recovered bioactives and extend their utilization across the agri-food sector, these extracts can be integrated into advanced preservation architectures, such as biomimetic materials engineered for sustainable fruit and vegetable supply chains (Tan et al., 2026). To provide a comprehensive overview, Table 1 systematically summarises these multiscale structural modifications and resulting structure–property relationships across hierarchical dimensions.

Importantly, these structure–property interactions extend beyond dead industrial matrices into living biological systems. Ultrasound-mediated mechanical perturbations trigger complex downstream cellular signalling events in tomato seeds. Acoustic exposure induces targeted microstructural alterations in the seed coat, enhancing water uptake and triggering metabolic activation via reactive oxygen species (ROS) signalling and oxidative enzyme pathways (Memiş et al., 2022; Nogueira et al., 2026). Emerging evidence further suggests that ultrasound-mediated stress responses can even provoke epigenetic modifications, inducing longer-term physiological adaptations and transgenerational plasticity across subsequent plant progenies (Farkas and Dobránszki, 2026).

6 Synergies, limitations, and future perspectives

Although acoustic and electro-physical operations are well characterised as standalone systems, their sequential or combined integration within a single cascade biorefinery layout remains an open research gap. Coupling cavitation-driven cell wall fragmentation with electroporation-mediated membrane permeabilisation provides a promising pathway for efficient, solvent-free process intensification.

However, several engineering bottlenecks restrict widespread industrial implementation. US process reproducibility is highly sensitive to acoustic energy attenuation, reactor geometries, and processing configurations, complicating cross-study standardisation and pilot-scale translation. PEF operations are inherently limited by the electrical conductivity and physical homogeneity of the tomato slurry, making them efficient for pumpable purees or juices but challenging for irregular solid matrices (Ghoshal, 2023). Additionally, excessive acoustic or electrical power inputs risk localized degradation of thermolabile molecules.

Future studies must focus on developing integrated, multiscale predictive models coupling cavitation physics, electroporation dynamics, and biological responses to accelerate the design of continuous, large-scale non-thermal platforms within sustainable circular bioeconomies.

7 Conclusion

Ultrasound and pulsed electric field technologies are powerful non-thermal methods that can modulate the structure and function of tomato systems across multiple hierarchical levels. Through distinct physical mechanisms, namely, acoustic cavitation and electroporation, both technologies selectively alter cellular boundaries, tissue organization, and mass transport phenomena. Quantitatively, these microstructural rearrangements translate into substantial macroscopic advantages, including enhanced carotenoid extraction yields, reduced industrial peeling resistance, and modified pulp rheology. Crucially, their operational efficiency is tightly constrained by the intrinsic biophysical properties of the matrix, where tissue porosity dictates acoustic propagation and electrical conductivity governs field uniformity, occasionally triggering synergistic localized thermal effects via Joule heating at high processing thresholds.

Beyond conventional extraction and preservation, emerging evidence indicates that these physical treatments can act as controlled elicitors in living tomato matrices, modulating biological signaling, physiological adaptations, and transgenerational stress responses. To secure the long-term stability and commercial application of these recovered bioactives, their integration into advanced architectures, such as sustainable biomimetic materials, offers a promising multidisciplinary path. Overall, connecting a multiscale structural framework with standardized analytical protocols, such as microscopical profiling and spectroscopy, provides a coherent platform to map structure–property relationships for next-generation sustainable processing technologies. Ultimately, future advancements in sustainable tomato processing will rely on predictive electro-fluid and mechanistic models capable of optimizing these synergistic non-thermal integrations within circular bioeconomies.

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Footnotes

Edited by: Andresa Gomes, University of São Paulo, Brazil

Reviewed by: Alkatuzzakia Akhi, Bangladesh Agricultural University, Bangladesh

Nanfeng Tan, Zhejiang University, China