FURTHER OBSERVATIONS ON JUVENILE AND ADULT HEDERA

Robbins , William J. (The New York Botanical Garden, New York, N. Y.) Further observations on juvenile and adult Hedera. Amer. Jour. Bot. 47(6) : 485–491. Illus. 1960.—Plants of arborescent Hedera helix sprayed with gibberellic acid produced juvenile shoots. Juvenile characters appeared in December to March from applications of gibberellic acid made from May to July. Gibberellic acid modified inflorescences toward a vegetative condition. Previous reports that seeds of arborescent Hedera helix produce juvenile plants were confirmed. Seedlings of a variant, Hedera helix ‘238th Street,‘ which has adult-shaped leaves on a vine type of growth produced vines with lobed leaves. Heavy pruning of arborescent Hedera helix caused the production of juvenile shoots.     

The Content of Elements of Mineral Nutrition in Leaf Parts of Different Colour in Plants of the variegata Type

The contents of some elements in green and whitish yellow parts of the leaves of Hedera helix and Acer negundo were examined. The contents of N, P and K were considerably higher in whitish yellow parts of the leaves compared with the green ones, and contrary to Ca in both plant species. Mg behaved like N, P and K in Hedera but like Ca in Acer. The same results were obtained in the leaves which were completely green or whitish yellow. The metabolic activities in different parts of the leaves were evaluated on the basis of the content of the individual elements. Differently coloured parts of the leaves of variegata type show differences not only in morphology and structure but also in their metabolic activity.     

Stomatal Responses to Light and Drought Stress in Variegated Leaves of Hedera helix

Direct and indirect mechanisms underlying the light response of stomata were studied in variegated leaves of the juvenile phase of Hedera helix L. Dose response curves of leaf conductance were measured with blue and red light in leaves kept in normal or in an inverted position. In the green portions of the leaves, the sensitivity to blue light was nearly 100 times higher than that to red light. No response to red light was observed in the white portions of the leaves up to 90 micromoles per square meter per second. Red light indirectly affected leaf conductance while blue light had a direct effect. Leaf conductance was found to be more sensitive to drought stress and showed a more persistent aftereffect in the white portions of the leaves. A differential effect of drought stress on the responses to blue and red light was also observed.     

Hepatoprotective and antioxidant effects of Hedera helix extract on acetaminophen induced oxidative stress and hepatotoxicity in mice

ABSTRACT

Exposure to high doses of acetaminophen is the most common cause of drug induced liver injury. We investigated the protective effects of Hedera helix extract against acetaminophen induced oxidative stress and hepatotoxicity using a mouse model. We used two control groups: group A was given 0.9% NaCl, group B was an acetaminophen control that was given a single injection of 600 mg/kg acetaminophen. T1?T4 groups were pretreated orally with different doses of H. helix extract. The mice were sacrificed and blood samples were collected to estimate the levels of glutathione peroxidase (GPx), malondialdehyde (MDA), superoxide dismutase (SOD) and total bilirubin. Liver samples also were used for histopathological studies. We found that acetaminophen significantly increased the levels of serum ALP, ALT, AST and MDA, and also significantly reduced the antioxidant factors, CAT, GPX and SOD. H. helix extract treatment produced a significant reduction in the levels of ALP, ALT, AST and MDA in serum and restored the levels of CAT, GPX and SOD to control levels. The histopathological findings were consistent with the biochemical findings. H. helix extract exhibited antioxidant and hepatoprotective effects against acetaminophen induced liver damage.     

The effect of light on the Og of plant tissues. I

Summary The increment of O_g in the guard cells in response to light as reported by previous investigators was found to be true qualitatively but not quantitatively.A special device was constructed, which permitted the elimination of the heat factor when the leaves were irradiated.In experiments 1 and 2 the relative values of the bell and the device were determined.In experiments 3 and 4 the relative effects of light were studied on younger and older leaves.In experiments 5 and 6 the effects were studied on old and young leaves ofHedera helix. This plant serves as a standard for comparisons of results obtained by subjecting plants to various factors of the environment.In experiments 7 and 8 the effect of light on young, middle-aged and old leaves ofHedera helix, when water was supplied, in one case but not in the other were studied. In experiment 8 the same leaf was treated in the device and then in the dark chamber, and once more in the light. The results are particularly enlightening.In the supplementary experiments, it was verified that the phenomenon of an increment of the guard cells and a decrement of the spongy parenchyma in O_g, when irradiated, is probably general.     

Light acclimation in leaves of the juvenile and adult life phases of ivy (Hedera helix)

Hoflacher, H. and Bauer, H. 1982. Light acclimation in leaves of the juvenile and adult life phases of ivy (Hedera helix). – Physiol. Plant. 56: 177–182. Light acclimation was investigated during the juvenile and adult life phases of the whole-plant-development in Hedera helix L. For this purpose, cuttings of the juvenile and adult parts of one single parent plant were grown under low-light (PAR 30–50 μmol photons m^?2 s^?1) and high-light (PAR 300–500 μmol m^?2 s^?1) conditions: CO₂ exchange, chloroplast functions, and specific anatomy of fully developed leaves differentiated under these conditions were determined. In juvenile plants the leaves formed under low and high light had light-saturated rates of net photosynthesis of 6.5 and 11.1 mg CO₂ (dm leaf area)^?2 h^?1, respectively. In adult plants the rates were 9.4 and 22.2 mg dm^?2 h^?1, indicating a more pronounced capacity for acclimation to strong light in the adult life phase. Higher photosynthetic capacities were accompanied by higher conductances for the CO₂ transfer through the stomata, leading to almost the same CO₂ concentration in the intercellular spaces. Thus, stomatal conductances were not primarily responsible for the different photo-synthetic capacities. The higher rates in adult and high-light grown leaves were mainly the result of formation of thicker leaves with more chloroplasts per unit leaf area. Expressed per chloroplast, the photosynthetic capacity, the Hill reaction, and the activity of ribulose bisphosphate carboxylase were almost identical in plants grown in low-light and high-light. Measurements of photosynthetic capacity and thickness of leaves of Hedera sampled from field habitats with contrasting light regimes confirm the results of growth chamber studies. It is, therefore, concluded that both life phases of Hedera are capable of acclimating to strong light, but that during the juvenile phase this capacity is not fully developed.     

THE BEHAVIOR OF TISSUE CULTURES FROM ENGLISH AND ALGERIAN IVY IN DIFFERENT GROWTH PHASES

In both the English and Algerian ivies, Hedera helix L. and Hedera canariensis Willd., leaf dimorphism of the juvenile and the mature, fruiting growth phases is pronounced, the former being a vine with lobed leaves and the latter a shrubby, upright form with entire leaves. Tissue cultures of English ivy started from stems of the two growth phases on the same plant consistently behaved differently, those from the juvenile stage invariably having the higher proliferation rate and larger cells. These differences were maintained consistently in monthly transfers over a period of two years. No medium was found which would support the growth of tissue subcultures of the adult stage of Algerian ivy, but all growth phases of the English ivy were cultured freely in a modified White's medium with additions of coconut water, naphthaleneacetic acid, enzymatic casein hydrolysate, and inositol. Cultures from individual open-pollinated seedlings of both species varied greatly in proliferation rate but were usually high.     

Distribution of Soluble 14C-Labelled Assimilates as Related to Distribution of 14C-Benzyladenine in Shoot Tips of Hedera helix

The relationship between benzyladenine and assimilate distributions in shoot tips of English Ivy (Hedera helix L.) was studied. Exogenous applications of benzyladenine resulted in small amounts of the hormone moving to the apical meristem. The large part of the recovered benzyladenine was found in the young leaves of the shoot tip. Benzyladenine increased the amount of ¹⁴C-labelled assimilates in the shoot tip, but the distribution pattern of those assimilates within the tip was like that of the untreated control.     

Photosynthesis in leaves of the juvenile and adult phase of ivy (Hedera helix)

Abstract Photosynthetic and anatomical parameters of leaves from the juvenile and adult part of an ivy plant (Hedera helix L.) have been determined and compared with each other. Light-saturated net photosynthesis (per unit leaf area) was about 1.5 times higher in adult leaves than in juvenile ones. The lower photosynthetic capacity of juvenile leaves was caused by a lower stomatal and especially a lower residual conductance to the CO₂-transfer. This corresponds with anatomical features of the leaves, i.e. lower stomatal frequency, fewer chloroplasts per cell, and – most important – thinner leaves, as well as with a less efficient photosynthetic apparatus measured as Hill reaction of isolated broken chloroplasts and activity of ribulose bisphosphate carboxylase. No differences in the respiration in light (relative to net photosynthesis) and in the CO₂-compensation concentration could be detected between the two leaf types. These observed anatomical and photosynthetic parameters of the juvenile and adult ivy leaves resemble those reported for shade and sun leaves, respectively, although the leaves investigated originated from the same light environment.     

Behavioural responses of the vine weevil, Otiorhynchus sulcatus, to semiochemicals from conspecifics, Otiorhynchus salicicola, and host plants

The vine weevil Otiorhynchus sulcatus is a parthenogenetic reproducing species which forages for suitable host plants at night, but is found congregated in dark places during the day. Frass of this weevil species is suspected to contain attractive compounds that are host‐plant related. Using a still‐air olfactometer, we tested adult vine weevils at night for their behavioural response to odours from conspecifics, feeding on a mixture of spindle tree (Euonymus fortunei) and yew (Taxus baccata), and to a sexually reproducing related species (Otiorhynchus salicicola), feeding on a mixture of ivy (Hedera helix) and cherry laurel (Prunus laurocerasus). Their attraction to conspecifics and O. salicicola appeared to be related to frass production. Freshly collected frass from O. sulcatus and from O. salicicola males and females was attractive. Prunus laurocerasus and H. helix have not been observed to be hosts of the vine weevil in the field. However, our tests showed that the vine weevil was attracted to mechanically damaged leaves of both plant species, whereas undamaged leaves were not attractive. Only undamaged young unfolding leaves of H. helix were also attractive. The attraction to odours from mechanically damaged host and non‐host plants suggested the involvement of compounds that are commonly found in many plant species. The involvement of plant compounds and/or aggregation pheromones in attraction to frass of the vine weevil and frass of the related weevil species O. salicicola is discussed.     

Water Pathways in Leaves of Hedera helix L. and Tradescantia virginiana L.

Hydraulic conductances of leaf tissues of Hedera helix and Tradescantiavirginiana leaves were measured. It was found that water couldflow most easily through the veins, but that the cell wallsof at least the ventral epidermis were more efficient at resupplyingwater lost from the epidermal tissue than was the mesophyllat rehydrating itself. Vein and bundle-sheath extensions, whichare characteristic of mesomorphic leaves (e.g. T. virginiana),seem to be important in maintaining a close hydraulic connectionbetween the epidermis and the vascular tissue. In leaves notcontaining vein and bundle-sheath extensions, typically xeromorphicleaves (e.g. H. helix), there is not such a close connectionbetween the epidermis and vascular tissue. This was shown inexperiments involving the sudden application of a reduced pressurepotential to either the epidermis or the other tissues of leaves,and the measurement of transient stomatal opening.     

Separation of Direct and Indirect Responses of Stomata to Light: Results from a Leaf Inversion Experiment at Constant Intercellular CO2 Molar Fraction

Previous work has shown that stomata respond directly to light,but it was not clear whether the only additional response isthrough CO₂, or whether some other metabolite is involved inthis response. Gas exchange experiments were done with normallypositioned and inverted leaves of Hedera helix to investigatethis problem. The macroscopic optical properties of the leavesand their anatomical structure were also studied. These experimentssnowed that there is no need to postulate the existence of amessenger other than CO₂ to explain the indirect response ofstomata to light. The experiments also showed that leaf inversionaffects both stomatal conductance and photosynthesis, and highlightthe difficulties involved in the interpretation of the effectof leaf inversion on stomata when stomatal conductance measurementsare not done concurrently with measurements of CO₂ flux densityand intercellular CO₂ molar fraction Key words: Hedera helix, ivy, gas exchange, leaf inversion, stomatal conductance, light, CO₂ flux density, photosynthesis     

Ribosomal RNA Gene Redundancy in Juvenile and Mature Ivy (Hedera helix)

Two-fold variation was found in rRNA gene redundancy in differentpopulations of ivy (Hedera helix). No consistent differencein rRNA gene number was observed between the juvenile and matureforms of the plant. Small significant differences between therRNA gene content of mature and juvenile forms and also betweenundeveloped buds and expanded leaves were found but these werein different directions in different populations.     

Phase Change in Hedera helix L: I. GIBBERELLIN-LIKE SUBSTANCES IN THE TWO GROWTH PHASES

Gibberellin-like substances were extracted from all parts ofthe juvenile and adult growth phases of Hedera helix. A numberof extraction techniques were employed and estimatos of theamount of gibberellin-like activity present were made usingthree bioassays. Gibberellin like substances were present inapproximately equal amounts in leaves at the two growth phasesand appeared to be localized, in the main, in chloroplasts.Apical ‘buds’ of the juvenile phase were shown tocontain higher levels of gibberellin-like substances than theiradult counterparts. Roots, which under normal circumstancesare associated only with the juvenile phase, contained highlevels of extractable gibberellin-like substances.     

Frost hardiness in the juvenile and adult life phase of ivy (Hedera helix L.)

The common ivy (Hedera helix L.) remains juvenileat its northern, eastern and altitudinal distribution limits although juvenileparts are largely killed by severe frost spells. In order to explain thisdiscrepancy we investigated the seasonal course of frost resistance in variousorgans of juvenile and adult parts of the same H. helixplants. Maximum frost resistance of leaves (LT₅₀–25°C) and axis (xylem parenchyma:LT₅₀ –29°C; cambium: LT₅₀–35°C) was quite the same in juvenile and adultparts. Thus, H. helix is able to acquire full frostresistance in its juvenile phase. However, hardening of leaves was slower anddehardening of axis was faster in juvenile parts. Leaves of juvenile partsremain 2 to 4 K less resistant than those of adult parts untilattaining the maximum resistance. This explains why mainly leaves of thejuvenile parts were damaged following severe frost episodes with temperaturesbelow –20°C. The occurrence of H.helix in its juvenile phase at the frost-caused distribution limitsmay be explained as follows: Leaves of juvenile plants may occasionally bekilled by severe frosts, but regeneration from dormant eyes enables survival.Loss of leaves may impede the change to the adult phase, but even if the plantsbecame adult frost killing of rest buds (2 to 3 K less resistantthan leaves) would induce rejuvenation.     

Function of leaf hamamelitol as a compatible solute during water stress treatment of Hedera helix L.

Hamamelitol is an unusual branched-chain sugar alcohol previously suggested to function as a leaf compatible solute. In this study, we have examined the leaf metabolism and intracelluiar compartmentalization of hamamelitol and other soluble sugars during long-term water stress treatment of Hedera helix (English ivy). Total leaf hamamelitol content was relatively low in greenhouse control plants, but increased 2-fold during water stress treatment to levels approaching those observed in field-grown plants (6–7 μmol g^?1 fresh weight). Using density gradient fractionation with non-aqueous solvents, we showed that hamamelitol occurs primarily in the cytoplasm and vacuoles of leaf mesophyll cells. During water stress treatment most of the increase in leaf hamamelitol occurred in the mesophyll cytoplasm, compensating osmotically for a decrease in cytoplasmic sucrose concentration. The maximum concentration of cytoplasmic hamamelitol was 155 mol m^?3 and occurred in field-grown plants. Labelling experiments showed that hamamelitol is slowly synthesized from ¹⁴CO₂ in leaves of H. helix, but is very long-lived (estimated t_1/2 of 4 years). Together, these data indicate that hamamelitol probably functions during long-term stress conditions as an osmotically active, compatible solute in plant leaves. We suggest that the signal for enhanced accumulation of hamamelitol during the water stress treatment was initiated by decreased plant growth and increased leaf sucrose hydrolysis.     

Phase Change in Hedera helix L: II. THE POSSIBLE BOLE OF ROOTS AS A SOURCE OF SHOOT GIBBERELLIN-LIKE SUBSTANCES

When synthesis was estimated by the agar diffusion techniqueboth basal and lateral adventitious roots of Hedera helix L.in the juvenile growth phase were shown to synthesize gibberellin-likesubstances. Seedlings and cuttings from juvenile ivy could begrown in water culture for several weeks. When roots were excisedfrom these plants shoot growth was reduced in comparison withthat of intact plants. The stems, apices, and leaves of derootedseedlings and cuttings contained lower levels of extractablegibberellin-like substances than comparable organs of intactplants. The major zone of gibberellin-like activity in intactplants co-chromatographed with gibberellins A₁ and A₃. Whenthese gibberellins were applied to plants grown in culture theywere found to promote growth of intact but not derooted plants.These findings are discussed in relation to the role of rootfactors and particularly gibberellins in phase change.     

Polyploidy and invasion of English ivy (Hedera spp., Araliaceae) in North American forests

Polyploidy is a common feature of agricultural weeds and natural area invaders. There are few studies comparing related diploid and polyploid exotics, however, and it is unclear what ecological and genetic factors favor the establishment of weedy polyploids. This research characterizes the geographic distribution and phenotypic characteristics of diploid Hedera helix and tetraploid Hedera hibernica, European species that are invading North American forests. To confirm the taxonomic affinity of invasive plants, we sequenced five non-coding cpDNA regions for 108 individuals (105 populations) as well as reference samples representing all species in the genus Hedera. Because diploid H. helix and tetraploid H. hibernica are poorly distinguished by morphology and DNA sequence, we used flow cytometry to determine their distribution (585 individuals). More than 90 % of sampled plants had cpDNA sequences identical or similar to H. helix sensu lato and H. hibernica. Diploid H. helix was dominant on the U.S. east coast (78.5 % of sampled plants) while tetraploid H. hibernica was dominant on the U.S. west coast (72.2 % of sampled plants), mirroring the species’ occurrence in maritime versus continental climates of Europe. Moreover, for sympatric occurrences in the Pacific Northwest, H. hibernica was larger and more frequently reproductive than H. helix. In a 2-year garden experiment, tetraploid H. hibernica had substantial architectural differences compared to diploid H. helix, including larger (but less numerous) leaves and thicker (but less branched) stems. Field experiments are needed to evaluate “pre-adaptation” (directional ecological filtering) and other factors mediating the invasion of H. helix and H. hibernica.     

Increase in CO2 concentration could alter the response of Hedera helix to climate change

Increasing CO₂ concentration ([CO₂]) is likely to affect future species distributions, in interaction with other climate change drivers. However, current modeling approaches still seldom consider interactions between climatic factors and the importance of these interactions therefore remains mostly unexplored. Here, we combined dendrochronological and modeling approaches to study the interactive effects of increasing [CO₂] and temperature on the distribution of one of the main European liana species, Hedera helix. We combined a classical continent‐wide species distribution modeling approach with a case study using H. helix and Quercus cerris tree rings, where we explored the long‐term influence of a variety of climate drivers, including increasing [CO₂], and their interactions, on secondary growth. Finally, we explored how our findings could influence the model predictions. Climate‐only model predictions showed a small decrease in habitat suitability for H. helix in Europe; however, this was accompanied by a strong shift in the distribution toward the north and east. Our growth ring data suggested that H. helix can benefit from high [CO₂] under warm conditions, more than its tree hosts, which showed a weaker response to [CO₂] coupled with higher cavitation risk under high temperature. Increasing [CO₂] might therefore offset the negative effects of high temperatures on H. helix, and we illustrate how this might translate into maintenance of H. helix in warmer areas. Our results highlight the need to consider carbon fertilization and interactions between climate variables in ecological modeling. Combining dendrochronological analyses with spatial distribution modeling may provide opportunities to refine predictions of how climate change will affect species distributions.