# Keep it cool and clean: Impact of home storage practices on fresh fruit and vegetable spoilage and efficacy of chemical versus green disinfectants

Damien Ballan, Jérôme Mounier, Dominique Amamana, Louis Coroller, Emmanuel Coton, Adeline Picot

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Damien Ballan, Jérôme Mounier, Dominique Amamana, Louis Coroller, Emmanuel Coton, et al.

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# Keep it cool and clean: Impact of home storage practices on fresh fruit and vegetable spoilage and efficacy of chemical versus green disinfectants

Damien Ballan, Jérôme Mounier, Dominique Amamana, Louis Coroller, Emmanuel Coton, Adeline Picot* ID

Univ Brest, INRAE, Laboratoire Universitaire de Biodiversité et Ecologie Microbienne, F-29280, Plouzané, France

<figure><img src="image_2.png" /></figure>

## ARTICLE INFO

**Keywords:**

Food waste

Consumer behavior

Cross contamination

Spoilage organisms

Food preservation

Citizen science

## ABSTRACT

Fresh fruit and vegetable (FFV) waste at consumer's level represents a considerable part of global waste in developed regions. Part of this waste stems from poor purchasing strategies and/or planning, but storage and hygiene practices may represent powerful means to reduce domestic FFV waste. The first aim of this study was to determine whether the hygiene status of storage compartments impacted microbial spoilage. Containers were inoculated with five spoilage organisms (3 molds and 2 bacteria), before storing FFV for 4 weeks at refrigerated or room temperature, and monitoring their deterioration. Temperature was the main factor influencing FFV loss. Contamination of surface compartments also significantly led to more spoilage on carrots and apples, but not on lemons and tomatoes. Different disinfectants were then tested to verify their efficacy against foodborne pathogens and spoilage organisms. After 5 min exposure, food contact disinfectant, sodium hypochlorite, white vinegar and rubbing alcohol were effective against all 12 tested organisms (3–4 log reduction) while lemon juice showed mitigated results. This is the first study assessing the link between storage contamination and FFV spoilage and the disinfection efficacy of so-called “Grandma’s remedies”. Overall, our results highlight the importance of consumer education about storage and cleaning/disinfecting practices.

## 1. Introduction

Food loss and waste have long been, and still remain, a burden in today's society as they lead to economic, social and ecological issues in both developed and developing countries. According to FAO definitions (FAO, 2011), food loss can occur throughout the food chain before reaching consumers, whereas food waste occurs at the retail and consumption level, both of which account for about 58 % of global food losses and waste (Vittuari et al., 2016). It is estimated that 40 % of consumers' food waste comes from fresh fruits and vegetables (FFV), making these products central to food waste reduction campaigns (FAO, 2011; Johnson et al., 2008). FFV loss along the food chain has been widely studied during the last decades (e.g. Alegbeleye et al., 2022; Balali et al., 2020; Karanth et al., 2023; Snyder & Worobo, 2018). However, studies primarily focus on pre-and post-harvest stages (e.g. cultivation, harvest, storage and distribution), as it is difficult to accurately measure FFV waste at the consumer level and determine the risk factors (Aschemann-Witzel et al., 2015). Domestic waste of FFV have two main sources: physiological decay, such as drying or over

maturation, or microbiological decay, mostly caused by molds and bacteria, although spoilage by protozoa or viruses can occasionally occur (Ballan et al., 2025; Pitt & Hocking, 2009; Samson et al., 2019; Snowdon, 1990).

In the earlier stages of the food chain, different methods are used to minimize the risk of losses, whether physiological or microbiological. These methods range from selecting resistant cultivars, applying phytosanitary treatments to prevent diseases in the field to using preservation technologies to either inactivate the microorganisms or inhibit their growth at post-harvest (e.g controlled atmosphere storage, cold storage, chemical or physical disinfection treatments) (Brecht et al., 2003; El-Ramady et al., 2015).

On the other hand, consumers often lack sufficient information when it comes to the best FFV storage and hygiene practices, usually relying on “common sense”, family habits or packaging instructions when available. For instance, most surveys related to storage practices underline sub-optimal usage of refrigerated compartments as well as excessive storage durations of perishable food products (Afriyie et al., 2023; Lagendijk et al., 2008; Terpstra et al., 2005). Indeed, knowledge

<footnote>* Corresponding author.</footnote>

<footnote>E-mail address: adeline.picot@univ-brest.fr (A. Picot).</footnote>
on optimal practices is lacking which can lead to malpractices and early losses of products (Marklinder et al., 2004). Besides, in everyday life, household members use a large array of surface disinfecting products, whether considered “natural” or “chemical” by consumers, to clean and/or disinfect FFV storage surfaces such as refrigerator drawers or fruit baskets (Truong, 2025). They can also use different products or methods to clean surfaces, but it is unclear which of them guarantee an effective disinfection. Besides, while many studies focused on foodborne pathogen contaminations in households, either on food itself or food contact surfaces, as it is essential to guarantee food safety at home (Evans & Redmond, 2019; Kennedy et al., 2005), only but few studies focused on FFV spoilage organisms.

Among consumer behaviors, storage and hygiene practices could thus represent powerful action levers to reduce FFV waste. Yet, to the best of our knowledge, no study has ever experimentally measured the impact of various storage and hygiene practices on FFV waste in order to provide consumers with best practices. In this context, the first main goal of this study was thus to accurately determine to which degree the contamination level of FFV storage compartments lead to FFV spoilage and waste, depending on the nature of FFV, the presence of surface wounds and the storage temperature (thereafter referred to as “spoilage dynamic assays”). The second objective of our study was to determine the disinfection efficacy of popular products used by consumers to clean storage compartments (e.g. fridge vegetable drawer or fruit basket) against both foodborne pathogens and common spoilage organisms (thereafter referred to as “disinfection assays”).

## 2. Material and methods

### 2.1. Spoilage dynamic assays

#### 2.1.1. Biological material

##### 2.1.1.1. Microbial strains.
Three molds (Penicillium expansum UBOCC-A-125030, Penicillium italicum UBOCC-A-125031, Botrytis cinerea UBOCC-A-125032) and two bacteria (Serratia marcescens UBOCC-A-325001 and Pseudomonas fluorescens group UBOCC-A-325002) were used for this experiment. All strains were isolated from spoiled fruits and vegetables collected in French households (Ballan et al., 2025).

#### 2.1.2. Experimental design

Four different fruits and vegetables were used during this experiment, namely tomatoes, lemons, carrots and “Ariane” variety apples. These products were selected as they are commonly consumed by French people, are available in retails during most of the year, and are sensitive to the selected organisms. Besides, these products were frequently thrown away by households throughout the year as observed in a previous study focused on the identification of microorganisms responsible for FFV waste at home (Ballan et al., 2025). The selected products were organically grown, to minimize the impact of phytosanitary product residues affecting the experiments.

Experiments were performed at either 7 °C (to mimic refrigerator conditions) or 19 °C (room temperature). Refrigerator temperature was selected considering previous results showing that the median temperature in 49 French household refrigerator FFV compartments was 7 °C (Ballan et al., unpublished results), while the room temperature was set at 19 °C following French government’s recommendation for optimal household temperatures (LegiFrance, 2022). Likewise, relative humidity was set at 80 % using a saturated solution of ammonium sulfate at 762 g/l according to the same previous results (Ballan et al., unpublished results). For each assay, both temperature and relative humidity were monitored using Thermo Hygro probes (Proges Plus, Lille, France).

Three levels of microbial load contamination of storage surfaces were evaluated. They corresponded to: i) a disinfected surface, ii) the median of storage surface contaminations in French households (10²

TFU/cm² molds and 10³/cm² CFU bacteria) and iii) the third quartile (10⁴ TFU/cm² molds and 10⁵ CFU/cm² bacteria) (Ballan et al., unpublished results) and were respectively designated as “none”, “moderate” and “strong” contamination levels.

Finally, the impact of wounds on spoilage was determined, using either unwounded or wounded products. Wounds were inflicted by cutting three scars (3-mm each) on the equator of the fruit or vegetable with a sterile scalpel, as described previously (Spadaro et al., 2011; Zhang et al., 2020).

In total, the experimental plan gathered 48 conditions (3 levels of microbial loads x 2 levels of storage temperatures x 2 levels of wounding x 4 types of FFV) of 10 FFV each. For logistical reasons (considering the large number of FFV to monitor), the entire experiment was conducted over three different dates, each corresponding to one level of microbial load.

Each experiment was conducted using a lidded hermetic plastic box in which FFV were placed for storage. For each level of microbial load x storage temperature x wounding, FFV were placed into two different boxes, each containing five FFV of each type. Beforehand, each plastic box was disinfected using 70 % ethanol then Duotex disinfectant (Laboratoires Rochex, Juvigny, France), and allowed to dry. Each FFV was cleaned with distilled water, before a 70 % ethanol bath for 10 min, then allowed to dry on sterile filters. The “wounded” fruits and vegetables then underwent cutting with a sterile scalpel. Finally, fruits from every condition were weighed, in order to compare their mass at the beginning and end of the experiment.

### 2.2. Inoculation methods

Fungal spore suspensions for contaminated conditions were obtained using 5-7-day-old cultures on malt and yeast extract agar (M2Lev) medium (20 g/l malt extract, 3 g/l yeast extract, 15 g/l agar). Conidia were collected using 2 ml of sterile water containing 0.01 % Tween 80 (Sigma Aldrich, Saint Quentin Fallavier, France) using a sterile L-shaped spreader. The obtained suspension was then collected and conidia numeration was performed using a Malassez hemocytometer. Dilutions were then performed using the same solution to reach the targeted spore concentration (between 1.0 and 5.0 10⁷ spores/ml).

For bacterial inoculation, fresh cultures on Plate Count Agar (PCA) medium (Difco, Saint-Ferréol, France) were used. One or two colonies were transferred into a tryptone salt sterile solution (Biokar, Allonne, France) and then vigorously vortexed, before optical density measurement at 600 nm. Bacterial concentration was calculated and the suspension underwent a series of dilutions in tryptone salt in order to reach the targeted concentration (between 1.0 and 5.010⁸ CFU/ml), before ongoing serial dilutions in tryptone salt solutions to reach the desired concentration. For both fungal and bacterial suspensions, one ml was inoculated and spread on the bottom of the plastic box using a sterile L-shaped spreader and allowed to dry for about 30 min under a laminar airflow. Control of inoculum size was performed by inoculating a mock box with the suspension which was sampled by swabbing once dried. Swab underwent vortexing for about 30 s in tryptone salt solution (9 g/l) then was diluted and plated on M2Lev medium supplemented with 5 mg/l penicillin G and 5 mg/l streptomycin (Sigma-Aldrich, Darmstadt, Germany) and PCA medium supplemented with 11 mg/l amphotericin B (Sigma-Aldrich, Darmstadt, Germany) to count fungi and bacteria, respectively.

Once the inoculum was dried, five fruits of each type (tomato, lemon, carrot and apple) were placed in each box as mentioned above and incubated according to the various conditions.

#### 2.2.1. Monitoring of spoilage dynamics

The experiment lasted 28 days, during the first 21 days, every 2-3 days, each plastic box was opened in sterile conditions to observe the products, take pictures and weigh the losses. After each observation, the products were randomly mixed in order to vary the fruits and vegetables
in contact with the contaminated surface at the bottom of the box. For each observation, fruit or vegetable presenting visible spoilage were noted as “damaged”. Of them, those with spoilage covering 50 % or more of the surface were noted as “spoiled”. Visible spoilage referred to the appearance of molds, browning, black spots, or rot on fruits and vegetables (Supplementary Fig. 1). They were then removed from the plastic box and weighed, as we considered that consumers would discard them in everyday life. Relative weight loss was also calculated following this formula:

$$ \text{Relative weight loss} = \frac{\text{Weight of discarded FFV}}{\text{Weight of discarded FFV} + \text{Weight of the remaining FFV}} $$

On days 7, 14, 21 and 28, each of the three experiments was subject to observation, making comparisons between conditions possible, on those specific dates.

### 2.2.2. Data treatment

Obtained data consisted of three main continuous dependent variables, namely number of “damaged” products, number of “spoiled” products and relative weight loss. The Area Under Disease Progression Curve (AUDPC) was calculated at day 7, 14, 21 and 28 for each of the three variables (de Mendiburu & Muhammad, 2020) following this formula:

$$ \text{AUDPC} = \sum_{i=1}^{n-1} \frac{y_i + y_{i+1}}{2} \times (t_{i+1} - t_i) $$

where $y_i$ is a measurement of “symptom” levels (number of damaged products, number of spoiled products or relative weight loss of considered product) at the i th observation, $t_i$ is the time (in days) at the i th observation and n is the total number of observations.

Analyses of variance (ANOVA) were performed to evaluate the effects of temperature, product type, contamination levels and presence of wounds on square root-transformed values of AUDPC for each of the three variables. If necessary, variables were aligned rank transformed to conduct nonparametric analyses of variance on factorial models using the ARTool package (Kay et al., 2025). In case of normal distribution of samples and homogeneity of variances (as validated by Shapiro and Levene tests), differences between conditions were then determined with multiple comparisons tests using Tukey HSD post-hoc test. If not the case, a Kruskal-Wallis test followed by a post-hoc Dunn's test was used. Statistical analysis and graphical representations were performed using RStudio V2022.07.2 + 576.

## 2.3. Disinfection assays

### 2.3.1. Biological material

Organisms used during this study belonged to 3 categories: pathogenic bacterial strains, namely *Staphylococcus aureus* ATCC 6538, *Listeria monocytogenes* ATCC 15313 and *Salmonella enterica* subsp. *enterica* serovar Typhimurium ATCC 13311, spoilage bacterial strains, namely *Pseudomonas fluorescens* group (UBOCC-A-325002), *Erwinia* sp. (UBOCC-A-325003), *Serratia marcescens* (UBOCC-A-325002), *Leuconostoc mesenteroides* (UBOCC-A-325004) and *Bacillus cereus sensu lato* (UBOCC-A-325005) and fungal spoilers, namely *Penicillium expansum* (UBOCC-A-125030), *Penicillium italicum* (UBOCC-A-125031), *Botrytis cinerea* (UBOCC-A-125032), *Cladosporium cladosporioides* complex (UBOCC-A-125033) and *Hanseniaspora uvarum* (UBOCC-A-224017). The pathogenic strains were those listed in the NF EN 13697 standard to determine the efficacy of disinfectants (AFNOR, 2023), while spoilers were isolated from spoiled FFV in French households from a previous study (Ballan et al., 2025).

#### 2.3.1.1. Products used for disinfection.
Five different products were tested on the different strains, namely 0.9 % sodium hypochlorite

(EuroTab, Saint-Just-Saint-Rambert, France), recommended by the French Agency for Food, Environmental and Occupational Health and Safety (ANSES) in their good practice guide (ANSES, 2013) for refrigerator disinfection; a commercial disinfectant compatible with food contact (0.3 % didecylmethylammonium chloride, Sanytol, Vaulx-en-Velin, France), 14° vinegar (St Marc, Massy, France), 60° rubbing alcohol (Paulette, Roche-Lez-Beaupré, France) and lemon juice (pH 1.92, Carrefour, Massy, France). The choice of these products was based on a survey conducted in 2023 on 2000 people representative of the French population (Azzib et al., unpublished results). The negative control corresponded to the use of sterile water.

### 2.3.2. Inoculation and disinfection protocol

The performed protocol was slightly modified from the ISO 13697 standard (AFNOR, 2023) to assess the efficacy of disinfection on contaminated food contact surfaces. For bacteria and yeasts, 1.0 to 5.0 10⁸ CFU/ml suspensions were prepared by resuspending fresh colonies, grown on PCA medium, in tryptone salt suspension. For molds, 50 µl of conidia suspensions with concentrations between 1 to 5.10⁷ spores/ml were prepared as described above. Then, 50 µl were deposited on the surface of a sterile empty Petri dish and allowed to dry for 1 h. The plastic from the Petri dish was made out of polypropylene which is a commonly-used type of plastic in storage compartments of domestic fridges (Ha & Kim, 2012). Once dried, 100 µl of the tested disinfecting product were added onto the spot for a contact duration of 5 min, before neutralization by adding 9.9 ml of a neutralizing solution (disodium phosphate 12 H₂O 100.80 g/l, sodium thiosulfate 5 H₂O 7.80 g/l, lecithin 3.00 g/l, L-histidine HCl H₂O 1.00 g/l, Tween 80 30.00 g/l, Humeau, Couëron, France) into the Petri dish and gently agitating the suspension to make sure all the microbial cells were resuspended. For *P. fluorescens*, *Erwinia* sp. and *L. mesenteroides*, contact with the disinfecting product was performed in liquid conditions (50 µl of the bacterial suspension and 100 µl of the disinfecting product, gently agitated) as these organisms did not tolerate drying onto the Petri dish. Dilutions were then plated using an “easySpiral Dilute” plating device (Inter-science, Puycapel, France) onto Tryptone Soya Agar (TSA) medium (Humeau, Couëron, France) followed by incubation at 30 °C for 3 days for bacteria enumeration or onto Malt Extract Agar (Humeau, Couëron, France) followed by incubation at 25 °C for 3 days for fungal organisms and enumerated using a Scan300 automatic colony enumerator (Inter-science, Puycapel, France). This method allowed for detection limits corresponding to a reduction of up to 4 log units for molds and 5 log units for bacteria and yeasts.

All tests were performed in triplicates. For each triplicate, the biocide efficacy was determined by calculating the log reduction obtained after biocide treatment as follows:

$$ \text{Log Reduction} = \log10(\text{Nw}) - \log10(\text{N}) \text{ where Nw is the mean cell concentration (in UFC/ml) of viable microorganisms after water exposition of the triplicate and N is the cell concentration of viable microorganisms after biocide exposition.} $$

## 3. Results

### 3.1. Spoilage dynamic assays

Fig. 1 shows the number of damaged FFV, showing visible spoilage, during a 28-day storage period at either 7 °C or 19 °C for each type of FFV, according to the contamination levels of storage surfaces (none, moderate or strong). Spoilage on FFV appeared gradually over time, without any drastic increase occurring within a short period of time (Fig. 1). As expected, refrigerated products usually took longer to present defects compared to those at room temperature. Overall, FFV stored at room temperature presented the most important damages, regardless of the considered FFV. Noteworthy, damages appeared at different rates depending on the considered FFV. For instance, apples and lemons revealed little to no damage during the course of the assay, even after 28
<figure><img src="image_3.png" /><figcaption>**Fig. 1.** Number of damaged FFV over time during storage at 7 °C (left panel) and 19 °C (right panel) for tomatoes (A and B), lemons (C and D), carrots (E and F), apples (G and H), and all FFV combined (I and J), previously wounded (dash lines) or not (solid lines) and according to the initial contamination levels of storage surfaces (none: no artificial contamination; moderate: $10^2$ TFU/cm² molds and $10^3$ CFU/cm² bacteria; strong: $10^4$ TFU/cm² molds and $10^5$ CFU/cm² bacteria).</figcaption></figure>

days, especially in refrigerated conditions (Fig. 1 C and G). However, carrots and tomatoes were more affected over time, to the point where all of them showed visible spoilage (as soon as day 12 for room temperature tomatoes and day 14 for room temperature carrots, Fig. 1 B and F). Most FFV defects corresponded to typical microbial spoilage from the inoculated microorganisms during the experiments: characteristic green mycelium of Penicillium spp. on apples and lemons, slimy spots from

bacteria or occasional Penicillium mycelium on carrots or grey mold on tomatoes from Botrytis cinerea for instance. Sporadically, other visual defects caused by microorganisms not inoculated during the test appeared throughout the experiments, such as observations of black mycelium on tomato peduncles or browning of a few parts of lemons.

For statistical data treatment, we calculated the AUDPC of three different variables: number of “damaged” products, number of “spoiled”

<caption>**Table 1**</caption>

P-values of analyses of variance showing the effects of the main factors (temperature, product type and contamination levels of storage area) and their two-way interactions on square-root transformed AUDPC values (sqrt of AUDPC) calculated according to the number of damaged products showing visible spoilage (N dam), the number of products showing visible spoilage >50 % (N spoiled) and relative weight loss (W loss) after 7, 14, 21 and 28 days of the experiment.

<table><thead><tr><th>Duration (days)</th><th>Variable (Sqrt of AUDPC)</th><th>Temp.</th><th>Prod.</th><th>Cont.</th><th>Temp. × Prod.</th><th>Temp. × Cont.</th><th>Prod. × Cont</th></tr></thead><tbody><tr><td rowspan="2">7</td><td>N dam</td><td>2.54 × 10<sup>-4</sup></td><td>0.099</td><td>0.270</td><td>0.723</td><td>0.685</td><td>9.34 × 10<sup>-3</sup></td></tr><tr><td>N dam</td><td>2.09 × 10<sup>-5</sup></td><td>1.73 × 10<sup>-2</sup></td><td>0.218</td><td>0.901</td><td>0.968</td><td>4.56 × 10<sup>-3</sup></td></tr><tr><td rowspan="2">14</td><td>N spoiled*</td><td>1.28 × 10<sup>-3</sup></td><td>0.140</td><td>0.871</td><td>0.378</td><td>0.079</td><td>2.33 × 10<sup>-3</sup></td></tr><tr><td>W loss*</td><td>3.71 × 10<sup>-4</sup></td><td>2.61 × 10<sup>-2</sup></td><td>0.732</td><td>0.469</td><td>0.117</td><td>1.18 × 10<sup>-3</sup></td></tr><tr><td rowspan="2">21</td><td>N dam*</td><td>1.49 × 10<sup>-13</sup></td><td>2.22 × 10<sup>-16</sup></td><td>6.04 × 10<sup>-4</sup></td><td>1.60 × 10<sup>-2</sup></td><td>0.298</td><td>1.06 × 10<sup>-5</sup></td></tr><tr><td>N spoiled*</td><td>1.27 × 10<sup>-9</sup></td><td>4.25 × 10<sup>-15</sup></td><td>1.18 × 10<sup>-2</sup></td><td>4.01 × 10<sup>-3</sup></td><td>0.216</td><td>5.16 × 10<sup>-6</sup></td></tr><tr><td rowspan="2">28</td><td>W loss*</td><td>8.61 × 10<sup>-9</sup></td><td>4.52 × 10<sup>-7</sup></td><td>0.065</td><td>0.080</td><td>0.322</td><td>9.87 × 10<sup>-7</sup></td></tr><tr><td>N dam*</td><td>4.41 × 10<sup>-14</sup></td><td>2.22 × 10<sup>-16</sup></td><td>0.207</td><td>1.57 × 10<sup>-2</sup></td><td>2.82 × 10<sup>-2</sup></td><td>2.19 × 10<sup>-5</sup></td></tr><tr><td rowspan="2"></td><td>N spoiled</td><td>8.16 × 10<sup>-10</sup></td><td>2.00 × 10<sup>-16</sup></td><td>0.434</td><td>0.320</td><td>0.156</td><td>5.23 × 10<sup>-8</sup></td></tr><tr><td>W loss</td><td>3.24 × 10<sup>-8</sup></td><td>8.34 × 10<sup>-10</sup></td><td>0.958</td><td>0.979</td><td>0.163</td><td>1.69 × 10<sup>-9</sup></td></tr></tbody></table>

Significant p-values appear in bold. Temp: Temperature of storage (7 °C or 19 °C); Prod: type of fruit or vegetable (Tomato, Lemon, Carrot or Apple); Conta.: Level of contamination (None, Moderate or High). Sqrt of AUDPC: Square root-transformed AUDPC values. Variables followed by a “*” symbol were aligned rank transformed to conduct nonparametric analyses of variance on factorial models (Kay et al., 2025). Apples were included from d21 since little to no damage was observed before. For the same reason, N Spoiled and W loss were included from d14.
products and relative weight loss from d7 to d28. Whatever the variable considered, the presence of wounds showed no significant effect, either alone or in two- or three-way interactions with the other considered factors (temperature, product type or contamination levels), this factor was thus discarded from further analyses (Supplementary Table 1). Considering the three remaining factors, the ANOVA revealed significant single effects of temperature or product type while the effects of contamination levels of storage surface were not significant (Table 1). Besides, significant two-way interactions were found, mostly between contamination of storage area with product type, and with temperature to a lesser extent (Table 1).

To facilitate multiple comparisons between conditions, we chose to display, in Fig. 2, the bar plot of square-root transformed AUDPC values, calculated according to the number of damaged products during the first 14 days of the experiment shown in Fig. 1. This variable allowed us to best discriminate between conditions, although similar conclusions could be drawn with the remaining variables (Supplementary Figs. 2 and 3). For each product considered, temperature had a significant effect on the AUDPC of damaged products (ANOVA with p-val = 0.0178; 0.0405; 0.0186; 0.0098 and $6.72 \times 10^{-6}$ respectively for tomatoes, lemons, carrots, apples and total products at day 14), confirming that at 19 °C, spoilage dynamics were faster and more severe. In addition, AUDPC of damaged carrots and apples also significantly varied according to contamination levels of storage area, in a temperature-dependent manner for apples (Fig. 2A and C). In other words, the levels of storage surface contaminations significantly impacted spoilage dynamics of apples only at 19 °C (p-val = 0.0228), while for carrots, the levels of contamination, whether moderate and/or strong, impacted spoilage for both temperatures (p-val = 0.0072 at 19 °C and 0.0409 at 7 °C).

In terms of waste measurement, it is worth mentioning that the mean relative weight loss for total products reached 12.3 % and 22.3 % at day 14 and 21, respectively in ambient conditions, while their refrigerated counterparts were 1.7 % and 9.5 %, respectively on the same dates.

## 3.2. Disinfection assays

Fig. 3 shows the biocidal efficacy of the five tested products, expressed as log reductions in microbial load relative to the water-treated control. Sodium hypochlorite, food contact disinfectant (Santol), rubbing alcohol and white vinegar showed a strong efficacy on all tested organisms, reaching the limit of detection of the protocol (about 3–4 logs for molds and about 4–5 logs for bacteria and yeasts), and thus corresponding to about 3–4 log reduction after a 5-min treatment. On the other hand, the efficacy of lemon juice showed more mitigated results depending on the considered organisms. A 5-min exposure of lemon juice was sufficient to yield a 4-log reduction for *Erwinia* sp., *L. monocytogenes*, *P. fluorescens* group and *S. marcescens*. However, lemon juice treatments showed mitigated results on *L. mesenteroides* and *S. aureus* (about 2–3 log reduction) and had little to no effect on *B. cereus*, *C. cladosporioides*, *H. uvarum*, *P. expansum*, *P. italicum* and *S. enterica* subsp. *enterica* serovar Typhimurium (about 0–1 log reduction).

## 4. Discussion

This study focused on evaluating the factors encountered during FFV storage in households that can lead to higher spoilage risks including storage temperature, the contamination level of FFV storage compartments, the presence of surface wounds on the products, the nature of FFV and the storage duration. As expected, storage temperature was the most prominent factor contributing to product spoilage, with significantly higher spoilage at room temperature regardless of the type of FFV. Indeed, higher temperature contributes to a higher metabolic and physiological activity in plant products, ranging from increased respiration, phytohormonal activity (ethylene and gibberellin production contributing to faster maturation among others), and water evaporation, which all contribute to shortened product shelf life (Moretti et al., 2010;

Vincent et al., 2020). Moreover, higher temperatures favor microorganism growth, with most spoilage organisms growing faster at room temperature as compared to refrigerated temperature (Johnson et al., 2008; Pitt & Hocking, 2009; Snowdon, 1990). In this context, the French Interprofessional Technical Center for Fruits and Vegetables (CTIFL) recommends optimal storage temperatures for domestic storage of different FFV, which are usually between 4 and 10 °C. A few exceptions can be noted, such as tomatoes as well as pineapples or bananas which can suffer from cold injuries and/or skin browning, when stored in refrigerators (CTIFL, 2023). Temperature itself must be considered alongside relative humidity, which has been described as an important factor for FFV preservation. Optimal hygrometry is usually of about 90–95 %, since lower values can lead to quicker dehydration of products and higher values favor microbial growth (Paull, 1999).

Although refrigeration has long been identified as a factor that extends the shelf life of FFV, consumers may prefer to store certain FFV at room temperature, whether to keep them visible and avoid forgetting to consume them, to make sure the product retains optimal organoleptic qualities as cold foods tend to reduce the perception of sweetness and aromas (Green & Nachtigal, 2012) or due to family habits. For example, the Waste and Resources Action Program (WRAP) reported that only 2–3 % of consumers in the UK considered that storing fruits and vegetables in fridges was important (Johnson et al., 2008) while Masson et al. (2017) observed that people usually lack knowledge about refrigerated compartments characteristics and advantages, such as temperature gradients in domestic fridges and the optimal position inside the refrigerator for different types of food. More recently, Ballan et al. (2025) showed that about three quarters of spoiled FFV collected in 49 households in the Brest area (Finistère, Brittany, France) were stored at room temperature. Therefore, educating consumers to make better use of refrigerators could reduce the amount of FFV waste caused by microbial spoilage. Thus, it is important to raise consumers' awareness on this subject, as it is already the case in many countries.

Secondly, we found that spoilage dynamics varied according to the type of FFV, apples and lemons showing more resistance, compared to more fragile products such as tomatoes. In particular, apples exhibited the lowest spoilage levels except under our worst-case conditions, that is at both ambient temperature and high contamination levels ($10^4–10^5$ CFU/TFU/cm²), suggesting that some products may be resilient to certain stresses up to a critical threshold. It is indeed well-established that some FFV are more rapidly spoiled than others and storage guidelines specific to a wide range of FFV are regularly made available (e.g., Johnson et al., 2008; Queensland government, 2023).

The level of contamination in storage surfaces also significantly contributed to FFV spoilage although this effect depended on the type of FFV and the storage temperature. At 7 °C, the levels of contamination on storage surfaces had no significant impact on the spoilage dynamics, except for carrots. At room temperature, significantly lower levels of apple and carrot spoilage were found on non-contaminated plastic boxes while the effect was non-significant for lemons and tomatoes. Low temperature is known to slow down microbial growth due to reduced metabolic activities, which may account for the lower impact of surface contamination on FFV spoilage. Conversely, at room temperature, the faster microbial growth may favour initial contamination from storage surface and subsequent colonization, although the extent of spoilage also depended on the considered FFV type. Such differences according to the FFV type is likely due to varying FFV defense mechanisms, intrinsic composition and physico-chemical parameters (e.g. pH). It should also be considered that we used a mixture of five microorganisms and therefore complex biological interactions, either antagonistic, neutral or synergistic, may occur. Surprisingly, we also observed during our assays that wounds (i.e. superficial scars) inflicted on products did not show any significant effect on tested FFV spoilage. In past studies, it was reported that other types of wounds, such as bruises or larger cuts, led to faster spoilage of products, either because of stress induced ripening or because it creates a vulnerable spot on products, suitable for microbial
<figure><img src="image_4.png" /><figcaption>**Fig. 2.** Barplot showing the mean square root-transformed AUDPC values of damaged products (visible spoilage) calculated after 14 days of experiment with standard deviations (n = 4). Overlaid points represent individual data: triangles correspond to wounded products, and circles to unwounded products. Different letters correspond to significant differences between groups for each level of temperature (p-val < 0.05), according to an ANOVA followed by Tukey HSD tests for carrots and Kruskal-Wallis followed by Dunn tests for apples.</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>**Fig. 3.** Dotplot showing the log reduction of spoilage fungi (in red), spoilage bacteria (in green) and human pathogenic bacteria (in blue) after a 5-min exposure with different products (n = 3).</figcaption></figure>

infections (Barth et al., 2009; Hyodo, 1991). Additionally, wounding may have triggered FFV physiological changes or localized defense responses, such as reactive oxygen species formation and plant cell wall strengthening with suberin and callose formation, which may have impacted microbial infection (Ackah et al., 2022; Gao et al., 2024; Lu et al., 2020). The differences in terms of the type of inflicted wound may account for the discrepancy between our study and the cited references. Overall, food spoilage is the result of complex biological and physico-chemical mechanisms resulting from both intrinsic and extrinsic biotic parameters (e.g. spoiler type, inter and intra-species interactions, FFV type and physiology) and abiotic parameters (e.g. FFV composition, pH, storage temperature and relative humidity).

Besides, it is also worth underlining that spoilage in uncontaminated conditions was observed although both fruits and storage surfaces were disinfected before the assay. In addition, sporadic microbial spoilage that did not correspond to any of the inoculated microorganisms during the test (black mold on tomatoes' peduncle or brownish dark spots on lemons). Some of this damage may be attributed to factors that were difficult to monitor in our experiment, such as the maturity stage of the products, presence of unnoticeable wounds on the product or contamination resulting from endophytes turning into spoilers when conditions were favourable (Gupta & Jawandha, 2010; Wenneker & Thomma, 2020).

Overall, these experiments revealed pioneering results about FFV storage in households, especially on the role of contamination of storage surfaces on the occurrence of microbial spoilage and the importance of the storage temperature and cleanliness of storage areas. The obtained results pave the way towards recommendations of best practices for consumers to guarantee a better preservation of their products. To go

further, it would be of interest to extend our study to a broader range of FFV, including more perishable categories such as berries or leafy vegetables, in order to pinpoint which are more prone to spoilage as well as to consider different types of wounds—such as impact damage from overloading—that may better reflect typical domestic storage conditions.

A second aspect of our study was to evaluate the hygiene practices of consumers by determining the disinfection efficacy of popular products against both foodborne pathogens, as required in biocide evaluation guidelines to ensure consumer's safety but also common spoilage organisms, which is usually much less taken into consideration. Yet, as we demonstrated, their presence on food contact surfaces can lead to increased spoilage and also some fungal spoilers may also produce mycotoxins with potent health risk to the consumer. The popular products we tested included sodium hypochlorite as well as a commercial disinfectant and alternative “green” products including rubbing alcohol, lemon juice and white vinegar, identified as the most used products by French consumers (Azzib et al., unpublished results).

Of all the tested products, the only one showing limited efficacy was lemon juice, and was particularly ineffective against *S. enterica* subsp. *enterica* serovar Typhimurium, and to a lesser extent against *S. aureus* which are both pathogenic organisms. Moreover, it showed limited efficacy against some of the studied spoilage organisms, especially molds. In contrast, white vinegar, also a weak acid, showed very good disinfection efficacies with up to 4 log reduction which is in agreement with previous results (Zinn & Bockmühl, 2020). Acetic acid has been described as an effective molecule since in acidic conditions, it is in an undissociated form ($pK_a$ = 4.76) and passively diffuses through cell membrane and affects the cytosolic pH homeostasis through its
dissociation into both acetate and H⁺ ions. Acetate contributes to both oxidative stress and turgor stress in cells while the H⁺ ions lower the cell pH and lead to severe metabolic disorders (Palma et al., 2018). As for lemon juice, the main acid, citric acid, showed poorer efficacy against microorganisms. This is due to citric acid's triprotic property combined with pKₐ values of 3.13, 4.76 and 6.40, making the molecule mostly undissociated at both acidic and neutral pH. Burel et al. (2021) revealed that citric acid was more effective against Gram-negative bacteria at pH of 9.5 compared to pH of 4.5 and 6.5. In the framework of this study, the pH of the tested lemon juice was really low (1.92), thus reducing its oxidative and biocidal properties. Different mechanisms of action were at stake for the other tested products. Sodium hypochlorite in bleach solution is known to disturb cell integrity. In solution, it dissociates into both hypochlorous ions (⁻OCl) and hypochlorous acid (HOCl), which have actions on key metabolic elements such as DNA, proteins, including enzymes, and amino-acids, but also on cell integrity (Jones & Joshi, 2021). Sodium hypochlorite is a widely used molecule in both industry and domestic contexts for its effective biocidal effect, ease of use and low cost, and remains one of the best choices when it comes to surface disinfection. However, the possibility of newly formed compounds, like chloramines that were reported potentially toxic (Wastensson & Eriksson, 2020), may explain why some consumers are reluctant to use it. The selected commercial food contact product (i.e. Sanytol) relies on the action of quaternary ammonium compounds whose action occurs mostly on the cell membrane, perturbing its polarity properties, leading to its solubilization and the liberation of cell contents. Quaternary ammonium compounds are highly effective disinfectants and are used in both industrial and domestic environments but they leave residues and should always be rinsed after disinfection (McBain et al., 2004).

Overall, the disinfection efficacy assay may also contribute to establishing efficient protocols for domestic cleaning and disinfecting of food contact surfaces, whether to eliminate pathogenic or spoilage microorganisms. Both sodium hypochlorite and Sanytol have attested efficacy against those organisms, with overall largely described biocidal effects on a large array of microorganisms (bacteria, molds, yeasts, viruses ...) (Gerba, 2015; Gerba & Kennedy, 2007; Morin et al., 2015); however, those products are sometimes negatively perceived by consumers which tend to consider industrial chemicals as harmful, whether for themselves or for the environment. To go further, future studies should also evaluate the efficacy of these disinfectants when a biofilm has formed in the fridge, for instance resulting from FFV decay. In any case, consumers must be reminded that disinfection alone does not guarantee an effective microbial removal, and a cleaning step to remove stains and debris should always precede disinfection.

## 5. Conclusion

This study showed that temperature was the main factor contributing to faster FFV spoilage. Contamination of surface compartments, investigated for the first time as a factor of fresh product spoilage in a domestic context, also significantly led to more spoilage on carrots and apples, but not on lemons and tomatoes. Disinfection methods were also investigated on both foodborne pathogenic bacteria and spoilage microorganisms, and showed that, with the exception of lemon juice, natural biocides (i.e. 14° vinegar and 60° rubbing alcohol) were as effective as commercial chemical disinfectants (i.e. sodium hypochlorite and quaternary ammonium) in the tested conditions. Overall, if part of FFV domestic spoilage roots from consumer supplying habits (overbuying, poor planning, compulsive purchases), post-purchase behaviors, including cold storage and cleaning practices, should also be considered as strong vectors to avoid FFV spoilage and thus food waste, raising the need for consumer education.

### CRediT authorship contribution statement

Damien Ballan: Writing – original draft, Visualization,

Methodology, Investigation, Formal analysis, Data curation, Conceptualization. **Jérôme Mounier:** Writing – review & editing, Validation, Supervision, Project administration, Methodology, Investigation, Funding acquisition, Conceptualization. **Dominique Amamana:** Methodology, Investigation. **Louis Coroller:** Writing – review & editing, Funding acquisition, Conceptualization. **Emmanuel Coton:** Writing – review & editing, Funding acquisition, Conceptualization. **Adeline Picot:** Writing – review & editing, Validation, Supervision, Methodology, Investigation, Funding acquisition, Conceptualization.

### Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

### Acknowledgements

This work was supported by the French National Research Agency (ANR) [grant numbers ANR-20-CE21-0006]

## Appendix A. Supplementary data

Supplementary data to this article can be found online at https://doi.org/10.1016/j.lwt.2025.118696.

### Data availability

Data will be made available on request.

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