Abstract
Water uptake is a critical issue for postharvest physiology of cut flowers. After harvest, cut flowers lose access to water from roots and sometimes develop emboli inside the xylem, which can disrupt water uptake and undermine flower longevity. The duration of dry storage (desiccation) before flowers are placed in a vase may affect rehydration capability. Despite the appreciated importance of desiccation time on water balance, the duration of desiccation that might cause irreversible damage in Lilium sp. L. is unknown. Therefore, we investigated effects of pre-vase dehydration on water uptake and postharvest quality of cut lilies. Stems of Lilium ‘Nashville’, ‘Santander’, and ‘Sorbonne’ were subjected to 0, 8, 24, or 48 hours of dehydration at 20 °C, then rehydrated in a solution containing 2% sucrose and a biocide. Water uptake in the first 24 hours of rehydration was significantly greater in dehydrated stems than in control (0 hour) stems. Although dehydration treatments increased water uptake in the first 24 hours after rehydration, total water uptake in dehydrated stems over the ensuing 9- to 10-day vase period was significantly less than in controls. In the vase, 48 hours of dehydration reduced the total water uptake in ‘Nashville’ by 27% and in ‘Sorbonne’ by 48%. Individual flower life and stem vase life were not affected significantly by dehydration treatment; however, ‘Sorbonne’ stems dehydrated for 24 or 48 hours had smaller flowers than controls and those that underwent the 8-hour dehydration treatment. ‘Nashville’ stems dehydrated for 24 hours showed visible leaf yellowing 3 days earlier than controls; ‘Sorbonne’ dehydrated for 48 hours showed leaf yellowing 2 days earlier. We conclude cut lilies have an ability to recover partially from significant dehydration and can restore water uptake initially, but pre-vase dehydration reduces total water uptake and affects some postharvest attributes negatively.
For most cut flowers, water balance is a vital factor for maximizing postharvest longevity. Unlike intact flowers, where life is ended either by color change, petal wilting, or abscission, cut flower life is often ended as a consequence of water stress, such as neck-bend of the peduncle, turgor loss of petals and leaves, and retarded flower opening (Van Doorn 1990). Water uptake of cut flowers is driven by transpiration. When water loss via stomata is more than water uptake, water stress ensues and reduces the longevity of cut flowers (Van Doorn 1997).
The factors affecting water uptake include xylem blockage, wounding-induced enzymes (Van Doorn 2012; Van Meeteren and Arévalo-Galarza 2009), and exudation of latex and other substances at the cut surface of the stem (Lineberger and Steponkus 1976). Xylem blockage may be induced by microbial growth (Aarts 1957; Van Doorn et al. 1989; Zagory and Reid 1986), aspirated air, and cavitation (Tyree and Sperry 1989) in the vessels resulting from cutting. The water column of the xylem vessels is held by tension. At the moment the stem is cut, the water column is broken and may result in embolism (Durkin 1980). The volume of air aspirated corresponds with the accessed volume of the vessel lumens opened by cutting (Van Doorn 1990).
During postharvest handling of cut flowers, dehydration stress can happen. Rosa hybrida L. cv. Cara Mia stems cut in air had reduced water uptake rates compared with stems cut under water (Durkin 1979). In R. hybrida cv. Sonia, leaf stomatal conductance decreased after ≥ 3 h of dehydration treatment. After 24 h of dehydration, there was a lower rate of water uptake and stems did not regain turgidity even after 36 h of rehydration (Van Doorn 1990). Another floral crop, chrysanthemum [Dendranthema grandifloram (Ramat.) Hemsl.] given prolonged dehydration time showed very low rates of water absorption (Durkin 1980). Joyce and Poole (1993) reported that stems of native Australian cut flowers (Verticordia plumosa or Verticordia nitens) failed to rehydrate after a 24- or 48-h dehydration treatment compared with dehydration of 6 or 12 h, which had no effect on longevity and flower abscission.
Anecdotally, lilies are thought to have an ability to “recover” after significant periods of dry storage. Lilium is one of the most important cut flower crops worldwide. Most hybrid lilies can be classified into five groups: Asiatic (A), Oriental (O), Longiflorum–Asiatic (LA), Longiflorum–Oriental (LO), and Oriental–Trumpet (OT) (Gill 2006). According to statistics released by U.S. Department of Agriculture, the wholesale domestic value of cut flowers was $295 million in 2020, with lilies accounting for $52.5 million (18%), greater than rose, gerbera, and chrysanthemum (U.S. Department of Agriculture, National Agricultural Statistics Service 2021).
To answer the question about the effects of dehydration on cut lilies, the purpose of this work was to determine whether pre-vase dry storage (dehydration) has detrimental effects on lily vase life. Using two O cultivars and one LA cultivar with four dehydration treatments, we tested the hypothesis that dehydration reduces subsequent water uptake and postharvest quality.
Materials and Methods
Plant material.
Dutch grown bulbs of Lilium ‘Nashville’ (LA), ‘Sorbonne’ (O), and ‘Santander’ (O) were planted in 60- × 40-cm crates with Lambert LM-111 planting mix (Lambert Peat Moss Co.; Rivière-Ouelle, Quebec, Canada) and grown in a glass greenhouse at Ithaca, NY, USA (23 °C day and 17 °C night) with 115 μmol⋅m–2⋅s–1 supplemental lighting by high-pressure sodium lamps from 7:00 AM to 8:00 PM. Stems with four buds were harvested when the first bud was puffy and colored.
Experimental design and treatments.
Stems were trimmed to 65 cm and leaves were removed from the bottom 15 cm of the stem before the dehydration treatments. Loose stems were dehydrated and placed flat on a table in a single layer in a growth chamber (20 °C, 10–12 μmol⋅m–2⋅s–1, 12 h of photosynthetic photon flux density, 60% to 70% relative humidity) for 0, 8, 24, or 48 h. After dehydration, the bottom 2 cm of the stem was trimmed in air and the were stems placed into test tubes with vase solution containing 2% sucrose (Van Doorn 2011) and 70 µL⋅L–1 commercial bleach for postharvest quality measurement (based on our initial trials). Each treatment had seven (‘Santander’) or 10 (‘Nashville’ and ‘Sorbonne’) replicates, with one stem per test tube (replicate) with a completely randomized design.
Measurements.
By weighing, solution uptake was calculated daily for the first 5 d, then each 2 d thereafter. The daily vase solution uptake was the test tube weight on the measuring date minus the weight on the previous date (W0) and was divided by days of uptake. Cumulative water uptake was calculated as the sum of all uptake determinations. Fresh weight changes were measured as follows: Stems were weighed on the date of harvest (FWo) then daily (FWa) thereafter. The fresh weight change was calculated as [(FWa – FWo)/FWo] × 100%. To measure flower life, individual flowers were tagged, and dates of opening and senescence recorded. Flower opening was defined as when the tips of the tepals separated, and stigma and anthers became visible. Senescence was defined as when the tepals lost turgor and became translucent or had marginal browning. Flower life was calculated as the difference of these dates. Vase life of a stem was calculated as the days between the first flower opening and the last flower senescing. Days to leaf yellowing was determined as follows: A leaf was defined to be “yellow” when ≥ 25% of its area was yellowed. The date of leaf yellowing was when at least three leaves per stem were yellow.
Statistical analysis.
Data were tested with one-way analysis of variance followed by the least significant difference test with P < 0.05 by using JMP Pro (version 15; SAS Institute, Inc., Cary, NC, USA).
Results
Dehydration.
Stems lost fresh weight during dehydration treatments, with longer durations causing more weight loss (Table 1). Weight loss in the 8-h dehydration treatment ranged from 2.2% to 5.1% of the initial fresh weight and increased to 9.6% to 13.8% in the 48-h treatment.
Dehydration effects on water uptake and fresh weight change in the first 24 h of rehydration of Lilium ‘Nashville’, ‘Sorbonne’, and ‘Santander’.
Fresh weight change in the first 24 h in the vase.
Cultivars varied in their ability to absorb water in the vase, in a manner dependent on prior dehydration treatment. In the first 24 hours of rehydration, ‘Nashville’ stems regained less fresh weight (as a percentage of initial weight) the longer they were dehydrated (Table 1). ‘Sorbonne’ dehydrated for 8 h hydrated readily in the first 24 h in the vase, less with 24 h of dehydration, and slightly lost weight as a result of 48-h dehydration treatment. Dehydration treatments had no effects on stem fresh weight gain in the first 24 h of hydration for ‘Santander’.
Water uptake in the first 24 h in the vase.
With ‘Nashville’, any dehydration pretreatment increased water uptake in the first 24 h in the vase (Table 1), but there was no difference between dehydration treatments. With ‘Sorbonne’, dehydration for 8 or 24 h increased the initial water uptake compared with 48 h of dehydration or nondehydrated controls. With ‘Santander’, any dehydration increased the initial water uptake, and uptake generally increased with longer dehydration (Table 1).
Time course of water uptake.
The three cultivars behaved similarly in that most water uptake happened in the first third (first 3 d) of the vase period (Fig. 1). After the first third of vase life, however, the cultivars differed significantly. ‘Nashville’ maintained water uptake throughout the vase period whereas ‘Santander’ had almost no additional water uptake after days 3 and 4. ‘Sorbonne’ was intermediate (Fig. 1).
Cumulative water uptake of Lilium. Data are means of 10, 10, and 7 stems for ‘Nashville’, ‘Sorbonne’, and ‘Santander’, respectively. Bars represent SE. If no bar is visible, it falls within the symbol dimension.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Cumulative water uptake of Lilium. Data are means of 10, 10, and 7 stems for ‘Nashville’, ‘Sorbonne’, and ‘Santander’, respectively. Bars represent SE. If no bar is visible, it falls within the symbol dimension.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Cumulative water uptake of Lilium. Data are means of 10, 10, and 7 stems for ‘Nashville’, ‘Sorbonne’, and ‘Santander’, respectively. Bars represent SE. If no bar is visible, it falls within the symbol dimension.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Total water uptake.
Longer dehydration pretreatments reduced total water uptake significantly in the vase (Table 1), with ‘Nashville’ and ‘Sorbonne’ showing a 20% to 48% reduction in total water uptake in the 48-h dehydration treatment compared with the controls. ‘Santander’ behaved differently; there was no difference in cumulative water uptake between the control and the 48-h dehydration treatment.
Stem fresh weight during the postharvest phase.
As with the time course of water uptake, cultivars presented broadly different patterns of weight change during postharvest, but in all three cultivars, stem fresh weight change tended to be similar between controls and stems dehydrated only 8 h (Fig. 2). With ‘Nashville’, controls and stems dehydrated 8 h continued to increase in fresh weight for at least 5 d, whereas the ‘Sorbonne’ and ‘Santander’ fresh weight increase was maximal by just 1 d. Thereafter, ‘Nashville’ stems dehydrated for 8 h maintained a positive fresh weight relative to the initial weight throughout the entire vase period, and those dehydrated 24 h or 48 h decreased by only ∼5% during the vase period. In dramatic contrast, controls and stems of ‘Sorbonne’ and ‘Santander’ dehydrated for any length of time lost 30% to 50% of their initial fresh weight during the vase period.
Fresh weight changes of Lilium. Data are means of 10, 10, and 7 stems for ‘Nashville’, ‘Sorbonne’, and ‘Santander’, respectively. Bars represent SE. If no bar is visible, it falls within the symbol dimension.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Fresh weight changes of Lilium. Data are means of 10, 10, and 7 stems for ‘Nashville’, ‘Sorbonne’, and ‘Santander’, respectively. Bars represent SE. If no bar is visible, it falls within the symbol dimension.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Fresh weight changes of Lilium. Data are means of 10, 10, and 7 stems for ‘Nashville’, ‘Sorbonne’, and ‘Santander’, respectively. Bars represent SE. If no bar is visible, it falls within the symbol dimension.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Flowering and flower life.
In ‘Sorbonne’, the large decrease in total water uptake in the 24-h and 48-h dehydration treatments inhibited flower opening, leading to smaller, malformed flowers (Fig. 3). There was, however, no effect of dehydration on stem vase life of ‘Sorbonne’ and ‘Santander’, and only the longest dehydration treatment reduced vase life compared with 8 h of dehydration in ‘Nashville’ (Table 2). Similarly, dehydration treatments had little consistent effect on individual flower life span (Table 3). However, after 48 h of dehydration, the third or fourth flower longevity of ‘Nashville’ and ‘Santander’ was reduced by ∼1 d compared with the control or 8-h dehydrated flowers. Dehydration accelerated leaf yellowing in ‘Nashville’ and ‘Sorbonne’, but had no consistent effect in ‘Santander’ (Table 2).
Appearance of Lilium ‘Sorbonne’ after 6 d in the vase after the indicated dehydration treatments.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Appearance of Lilium ‘Sorbonne’ after 6 d in the vase after the indicated dehydration treatments.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Appearance of Lilium ‘Sorbonne’ after 6 d in the vase after the indicated dehydration treatments.
Citation: HortScience 57, 11; 10.21273/HORTSCI16766-22
Dehydration effects on vase life and days to leaf yellowing of Lilium ‘Nashville’, ‘Sorbonne’, and ‘Santander’.
Effects of dehydration longevity on individual flower life by flower position of Lilium ‘Nashville’, ‘Sorbonne’, and ‘Santander’.
Discussion
Lilies subjected to dehydration treatments were able to regain water uptake and fresh weight after 24 h of rehydration. However, longer dehydration treatments reduced water uptake and led to more rapid fresh weight loss. Longer dehydration durations lead to smaller and malformed flowers. Dehydration duration had little consistent effect on individual flower life in three cultivars.
Van Doorn (1990) found that air is aspirated into xylem elements very rapidly when plant parts are cut in air. In most species, aspiration stops after 30 min, and little additional air enters the stem. Van Meeteren et al. (2006) demonstrated that aspirated air was mainly responsible for initial xylem occlusion. Furthermore, as air exposure time increases, cell wall wettability decreases, which makes entry of water into the lumen of the xylem conduits opened by cutting more difficult (Van Doorn and Otma 1995). Fanourakis et al. (2021) also explained that decreased transpiration of cut chrysanthemum during dry handling was a result of stomatal closure.
Our results are consistent with these findings. In the first 24 h after rehydration, lilies subjected to dehydration restored water uptake and gained more fresh weight than nondehydrated stems (Table 1, Figs. 1 and 2). This possibly suggests lilies have emboli repair mechanisms. A relatively shorter dehydration time of 8 h increased water uptake compared with controls in ‘Nashville’ and ‘Santander’. Similarly, lisianthus (Eustoma gradiflorum Salisb.) had more water uptake in the first 24 h in the vase after 1 week of 2 °C dry storage compared with wet storage (Ahmad et al. 2012). Also, gladiolus (Gladiolus L.) subjected to dry storage for up to 36 h at 5 °C had rapid water uptake and recovery of fresh weight with just 1 h of rehydration (Costa et al. 2017). Compared with gladiolus (Costa et al. 2017) and lily, chrysanthemum stems decreased uptake and had limited water flow after holding in air for more than 2 h, and could not restore water flow after flower stems were placed in water (Van Meeteren et al. 2006). These studies show different floral crops have different tolerances to dry storage. For lily, longer dehydration periods (e.g., 48 h) reduced the ability of stems to absorb water during the ensuing vase period (Table 1), but 8 h of dehydration did not show obviously negative effects. Although lily stems restored water uptake and fresh weight in the first 24 h in the vase, the dehydration still reduced cumulative water uptake by 7% to 59%, depending on the cultivar and desiccation duration (Table 2, Fig. 3). Woltering and Paillart (2018) also pointed out that cut roses stored dry for 2.3 d at 6 °C or 28 d at 0.5 °C had less water uptake compared with nonstored stems as a result of reduced stomatal functionality.
With ‘Sorbonne’, stems in any dehydration treatment had inferior postharvest qualities, including smaller opened flowers (Fig. 3) and early development of leaf chlorosis (Table 2). Similarly, Capsicum annuum L. ‘Rio Light Orange’ and ‘Cappa Round Red’ cut stems had more wilted foliage if subjected to dry storage for 1 week vs. storage in water (De F. M. França et al. 2017). Rose was also not tolerant to dehydration, which causes less water uptake and inferior postharvest quality in the vase phase (Ahmad et al. 2014; Cevallos and Reid 2001; Van Doorn and Suiro 1996). Daffodil (Narcissus L.) also had more than a 50% shorter vase life in dry storage at 12.5 °C vs. wet storage (Cevallos and Reid 2001). Woltering and Paillart (2018) indicated that decreased stomatal functionality was involved in poor flower performance parameters, such as vase life and flower weight changes.
As with Eremurus (Ahmad et al. 2014), Lilium cultivars differed in desiccation tolerance. Elibox and Umaharan (2010) also showed that vase life correlated positively to daily water uptake rates during the vase period, and the cultivars that absorbed more water exhibited longer vase life and delayed the symptoms of water stress. The factors affecting water uptake may include transpiration rate (Van Doorn and Reid 1995) and air emboli in xylem vessels (Van Ieperen et al. 2002). Van Doorn and Reid (1995) reported cultivars with lower transpiration rates had slower development of vascular occlusion after exposure in the air compared with other cultivars. In our study, ‘Nashville’ and ‘Santander’ showed greater overall tolerance to dehydration than ‘Sorbonne’, agreeing well with the previously cited work. It seems likely that significant cultivar differences can be expected among the many commercially important lily cultivars and groups.
Our results therefore suggest that lilies should not be held dry at 20 °C for more than 24 h after harvest. Theoretically, if cut lilies were exposed to lower temperature, the critical exposure time could be longer because of a lower transpiration rate, with less water loss. Correspondingly, Xue et al. (2019, 2020) presented that dry storage at 0 to 4 °C improved cut peony (Paeonia lactiflora Pall.) vase quality compared with wet storage by accelerating carbohydrate metabolism and increasing water uptake efficiency. We found that the reduction of total water uptake in cut lilies resulting from long-term dehydration reduced the diameter of opened flowers and time to leaf yellowing, but flower life in general was unaffected. For further research, the measurement of hydraulic conductance and comparison of xylem anatomy among cultivars might be useful to provide more supporting evidence for dehydration stress on cut lilies. Our research findings demonstrated that cut lilies are tolerant to a water deficit, and that cultivars have a varied response to dehydration. In general, more than 24 h of nonrefrigerated dehydration is harmful to water uptake and flower opening of cut lilies.
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