## 1 Introduction  
High-voltage pulsed electric fields (PEFs) can induce the electrical breakdown and rupturing of cell membranes in biological tissues [1]. PEF treatment transiently increases the plasma membrane permeability by forming nanopores, thereby facilitating the transport of otherwise cell-impermeant molecules, which increases electrical conductivity. This phenomenon is referred to as electroporation or electro-permeabilization [2, 3, 4]. If the cell succeeds in restoring homeostasis by repairing its plasma membrane, the effect is transient and termed reversible electroporation (RE). However, when the PEF pulse parameters exceed certain thresholds, irreparable damage occurs in the plasma membrane, resulting in cell death. This effect is termed irreversible electroporation (IRE) [5]. A clinical application of RE and IRE is minimally invasive tumour treatment [6, 7, 8, 9, 10], with IRE being particularly effective for tumours near critical blood vessels in the liver, pancreas, and brain due to its non-thermal mechanism that preserves vascular structures [11]. Commonly, series of unipolar square wave pulses [12], characterized by parameters such as the number of pulses (NP), pulse voltage (PV), pulse width (PW), and pulse interval (PI) (see Figure 1d for parameter definitions), are locally applied to the treated tissue over a needle applicator [13]. The distance between the straight needle electrodes and the PV determines the maximum applied PEF field strength [14]. Numerous studies have explored the effects of PEF parameters across in vitro [15; 16], in vivo [17; 18], and in plant-based models [19; 20]. Towards higher NP, the PV, PW, and smaller PI favour IRE over RE [5; 21; 22]. Eventually, if the PEFs start to cause Joule heating, thermal ablation will also contribute [14; 23]. However, thresholds between the regimes may not only depend on the above-mentioned pulse parameters, but also on the tissue type [24] and shape of the needle applicator [21; 25; 26]. Understanding these influences is albeit crucial with respect to patient-specific individual treatment planning [27]. A study in rabbit liver by [PERSON] et al. found thresholds for RE and IRE at applied electric fields of 460 and 700 V/cm, respectively (NP = 8, PW = 100 \(\upmu\)s, PI = 1 s) [28]. However, for RE treatment in humans, eight electric pulses with a PW of 100 \(\upmu\)s are commonly delivered at an electric field of 1000 V/cm or higher [29]. An optimization study on clinical IRE treatments using a NanoKnife on swine liver found that increasing PW beyond 50 \(\upmu\)s and NP beyond 50 did not significantly increase the ablation size due to IRE [30]. Furthermore, PW > 70 \(\upmu\)s and NP > 70 caused white necrotic tissue next to the electrode because of thermal coagulation, similar to that seen in radiofrequency and microwave ablation [30]. It should be noted that plant tissues seem to require higher PW of around 100 \(\upmu\)s-1000 \(\upmu\)s for successful PEF treatment [31]. Importantly, pulse parameter experiments necessitate a substantial quantity of viable tissue, which can be either living animal or plant tissue. PEF effects, especially IRE and RE, were also explored at the single-cell level, but tissues present a more complex scenario  
Figure 1: **(a)** EIS measurement system featuring an open-ended coaxial four-point probe (abbreviations W, WS, R, and C correspond to working, working sense, reference, and counter electrodes, respectively). The excitation voltage was applied between electrodes W and C, while impedance spectra were acquired between WS and R. **(b)** Two-needle PEF treatment system. **(c)** Anatomical regions within potato tissue. **(d)** Definition of the unipolar pulse parameters (abbreviations NP, PV, PW, and PI correspond to number of pulses, pulse voltage, pulse width, and pulse interval, respectively). due to the proximity of cells and the inherent tissue inhomogeneity [32]. In contrast to animal models, metabolic activity, including cellular respiration, persists in plant models even after harvesting [33].