Early responses of drought-resistant and -susceptible tomato plants subjected to water stress

Three-week-old seedlings of one drought-susceptible tomato cultivar (Lycopersicon esculentum cv. New Yorker) and two drought-resistant species of tomato (Solanum pennellii andLycopersicon chilense) were subjected to various degrees of PEG 8000-induced water stress from –0.017 to –1.0 MPa for a duration of 24 h so that their early responses to water stress could be compared. Such a comparison would determine if there was a relationship to root cytokinin levels following sudden induction of water stress in the drought-resistant species. Transpiration rates of leaves were monitored throughout the 24-h period, shoots were evaluated for leaf water potential (LWP), and roots were extracted for levels oft-zeatin riboside (t-ZR) and dihydrozeatin riboside (DHZR) using a monoclonal antibody enzyme immunoassay. Transpiration rates were evaluated gravimetrically by difference every 6 h up to 24 h. Transpiration rate decreased with increasing PEG levels and passage of time in all three species, measured at 6 and 12 h, logarithmically in the case of the twoLycopersicon species and linearly in the case ofSolanum. From 12–18 h (while plants were in darkness), transpiration rate was a function of the level of PEG only and not time in all three species. When light resumed from 18–24 h, only 5.pennellii showed no further decrease in transpiration rate over time with increasing PEG. Drought-susceptibleL. esculentum had a stronger linear decrease in LWP with increasing PEG 8000 concentration than the other two species.L. esculentum also had a higher initial transpiration rate than did either of the drought-resistant species. The two drought-resistant species showed less change in LWP with 5.pennellii having a small decrease andL. chilense having little change. OnlyS. pennellii exhibited a decrease in roott-ZR levels, which may imply a role for root cytokinin within the first 24-h exposure to water stress in this species.L. esculentum exhibited no change in roott-ZR. The levels oft-ZR inL. chilense were less than that ofL. esculentum but showed only a slight decrease with increasing PEG.S. pennellii andL. chilense, although both drought-resistant tomato species, showed different patterns of response with respect to pattern of decline in transpiration rate, LWP, and roott-ZR levels.     

Transpiration in wet meadows and some other types of Grassland

Transpiration rate in four types of alluvial meadows has been studied. Water wapour output of different grassland communities seems to be proportional to their biomass, bearing no relation to the species composition even though transpiration of individual species differed considerably. The relation between the above-ground biomass and the daily transpiration has been ascertained for wet, mesophytic and dry grassland and seems to be constant under similar sets of environmental conditions. The coefficient for estimating the daily transpiration in meadows from their biomass is presented.     

Effects of wind velocity on stomatal conductance and consequences of leaf rolling on water uptake in tall fescue

Leaf form and stomatal characteristics play an important part in the strategy of tall fescue (Festuca arundinacea Schreb) to limit water losses. Transpiration rates of artificially rolled leaves measured by potometer at varying wind velocities support this assertion.     

Transpiration in seven plant species colonizing a fishpond shore

In 7 species (Eleocharis palustris R. Br.,Juncus bufonius L.,Gypsophila muralis L.,Trifolium repens L.,Agrostis stolonifera L.,Potentilla anserina L. andAchillea millefolium L.) growing in a gradient of habitats from aquatic to terrestrial, on a sandy fishpond shore in Southern Bohemia, Czechoslovakia, the daily course of transpiration rate and water content was assessed gravimetrically in their cut-off transpiring parts on two typical summer days. Transpiration rate was largely controlled by microclimate but depressions occurred inJuncus and in some species growing in the drier habitats. The highest instantaneous transpiration rate and total daily transpiration per unit dry weight, about 100 mg g^?1 min^?1 and 50 g g^?1 d^?1, respectively, were recorded inAchillea andTrifolium. The water turnover rate ranged from 16.1 g g^?1 (H₂O) d^?1 inJuncus to 10.7 inGypsophila. The estimated daily maximum water saturation deficit was highest inEleocharis (45.7%) and lowest inAchillea (15.0%). The adaptational significance of the observed phenomena is discussed.     

Osmotic adjustment in leaves ofLycopersicon esculentum andL. pennellii in response to saline water irrigation

Two tomato species (Lycopersicon esculentum andL. pennellii) were grown under unheated plastic greenhouse and irrigated with 0 or 140 mM NaCl. Salinity induces a more important reduction in predawn leaf water potential (ψ_pd) inL. esculentum than inL. pennellii. In both species the osmotic adjustment was achieved by active solute accumulation. The leaf water potential at turgor loss point (ψ_tlp) seemed to be controlled by leaf osmotic potential (ψ_os). The results revealed the existence of limits to the accumulation of osmotic solutes in leaf tissues and the existence of an ontogenetic effect on the solute accumulation. In both species, but essentially inL. pennellii the inorganic solutes contribution especially Na⁺ and Cl^? accumulation to ψ_os was higher than the organic solutes. Therefore, wild species save energy more markedly.     

Osmotic adjustment in Lycopersicon esculentum and L. Pennellii under NaCl and polyethylene glycol 6000 iso–osmotic stresses

Changes in leaf solute contents in response to saline (NaCl) and osmotic (polyethylene glycol, PEG, 6000) stresses were measured in three different salt tolerant cultivars of Lycopersicon esculentum (L.) Mill. (Pera, P-73 and Volgogradskij), and its wild relative L. pennellii (Correll) D'Arcy accession PE-47. Iso-osmotic stresses (–0. 5 MPa) of NaCl (140 mM) and PEG 6000 (150 g l^-1) were applied to one-month old plants for 3 weeks. Decreasing leaf dry weight was similar in L. pennellii or L. esculentum cv. P-73 and Volgogradskij under both stresses, while leaf dry weight of L. esculentum cv. Pera decreased more under PEG stress than under NaCl stress. Water contents decreased in all the PEG treated populations, while their calculated solute potential (Ψ_s increased. Under osmotic stress, the total ion contents decreased in relation to control, whereas organic solutes (sugars, amino acids and organic acids) markedly increased in both tomato species, specially in the tomato cultivars, where these solutes represented 50% of the Ψ₅ calculated. Soluble sugar increase was three times higher in leaves of L. esculentum than in the leaves of L. pennellii. Free proline increased under both stresses and its content was highest in L. esculentum and in L. pennellii, respectively, under NaCl and PEG stresses. Nevertheless, the contribution of this metabolite to Ψ_s did not exceed 5%, irrespective of treatment and species. The greater organic solute accumulation in L. esculentum than in L. pennellii– which was not reflected in their Ψ₅ values – was not correlated with the tolerances of the two species to osmotic stress. Therefore, osmotic adjustment may not be the only process influencing salt and drought tolerances in tomato; the ability of plants to regulate their metabolic and physiological functions could also play an important role under these harmful conditions. The possible roles of inorganic solutes and metabolites in osmotic adjustment, energetic metabolism and redox regulation are discussed     

Hydraulic adjustments in aspen (Populus tremuloides) seedlings following defoliation involve root and leaf aquaporins

Main conclusion

Changes in root and leaf hydraulic properties and stimulation of transpiration rates that were initially triggered by defoliation were accompanied by corresponding changes in leaf and root aquaporin expression. Aspen (Populus tremuloides) seedlings were subjected to defoliation treatments by removing 50, 75 % or all of the leaves. Root hydraulic conductivity (L_pr) was sharply reduced in plants defoliated for 1 day and 1 week. The decrease in L _pr could not be prevented by stem girdling and it was accompanied in one-day-defoliated plants by a large decrease in the root expression of PIP1,2 aquaporin and an over twofold decrease in hydraulic conductivity of root cortical cells (L _pc). Contrary to L _pr and L _pc, 50 and 75 % defoliation treatments profoundly increased leaf lamina conductance (K _lam) after 1 day and this increase was similar in magnitude for both defoliation treatments. Transpiration rates (E) rapidly declined after the removal of 75 % of leaves. However, E increased by over twofold in defoliated plants after 1 day and the increases in E and K _lam were accompanied by five- and tenfold increases in the leaf expression of PIP2;4 in 50 and 75 % defoliation treatments, respectively. Defoliation treatments also stimulated net photosynthesis after 1 day and 3 weeks, although the increase was not as high as E. Leaf water potentials remained relatively stable following defoliation with the exception of a small decrease 1 day after defoliation which suggests that root water transport did not initially keep pace with the increased transpirational water loss. The results demonstrate the importance of root and leaf hydraulic properties in plant responses to defoliation and point to the involvement of PIP aquaporins in the early events following the loss of leaves.     

Unilateral incongruity in crosses involvingLycopersicon pennellii andL. esculentum is distinct from self-incompatibility in expression,timing and location

Both interspecific and intraspecific mechanisms restrict the exchange of genes between plants. Much research has focused on self incompatibility (SI), an intraspecific barrier, but research on interspecific barriers lags behind. We are using crosses betweenLycopersicon esculentum andL. pennellii as a model with which to study interspecific crossing barriers. The crossL. esculentum×L. pennellii is successful, but the reciprocal cross fails. Since the cross can be successfully made in one direction but not the other, gross genomic imbalance or chromosomal abnormality are precluded as causes. We showed that the lack of seed set observed in the crossL. pennellii×L. esculentum is due to the inability of pollen tubes to grow more than 2–3 mm into the style, whereas S1 crosses show continued slow pollen tube growth but, also, fail to set seed. These results indicate that the unilateral response is a barrier distinct from SI, differing from SI in the timing and location of expression in the style. We therefore suggest that this unilateral response in theL. pennellii×L. esculentum cross is more accurately referred to as unilateral incongruity (UI) rather than interspecific incompatibility. Periclinal chimeras were used to determine the tissues involved in UI. The results of crosses with the available chimeras indicate that the female parent must beL. pennellii at either LI (layer 1) or both LI and LII (layer 2) and the male parent must beL. esculentum at either LII or both LI and LII to observe UI similar to that seen in theL. pennellii×L. esculentum cross. Pollinations with a mixture of pollen fromL. pennellii and from transgenicL. esculentum plants harboring a pollen-specific GUS reporter gene marker were used to ascertain whether the growth of the pollen tubes of either species was modified as a possible means of overcoming UI. We found no evidence of communication between the two types of pollen tubes to either enhance or restrict all pollen tube growth.     

Mitochondrial DNA Sequence Divergence among Lycopersicon and Related Solanum Species

Sequence divergence among the mitochondrial (mt) DNA of nine Lycopersicon and two closely related Solanum species was estimated using the shared fragment method. A portion of each mt genome was highlighted by probing total DNA with a series of plasmid clones containing mt-specific DNA fragments from Lycopersicon pennellii. A total of 660 fragments were compared. As calculated by the shared fragment method, sequence divergence among the mtDNAs ranged from 0.4% for the L. esculentum-L. esculentum var. cerasiforme pair to 2.7% for the Solanum rickii-L. pimpinellifolium and L. cheesmanii-L. chilense pairs. The mtDNA divergence is higher than that reported for Lycopersicon chloroplast (cp) DNA, which indicates that the DNAs of the two plant organelles are evolving at different rates. The percentages of shared fragments were used to construct a phenogram that illustrates the present-day relationships of the mtDNAs. The mtDNA-derived phenogram places L. hirsutum closer to L. esculentum than taxonomic and cpDNA comparisons. Further, the recent assignment of L. pennellii to the genus Lycopersicon is supported by the mtDNA analysis.     

Stomatal Response to Light of Solanum pennellii, Lycopersicon esculentum, and a Graft-induced Chimera

To learn how species differences in stomatal behavior are regulated, the response of epidermal and leaf diffusive resistance to light was investigated in Lycopersicon esculentum Mill., Solanum pennellii Corr., and a periclinal chimera having an S. pennellii epidermis and an L. esculentum mesophyll that was produced from a graft of the two species. S. pennellii has about 23% fewer stomata per square millimeter than does L. esculentum, and the two species have contrasting stomatal sensitivities to light. The abaxial stomata of L. esculentum open in dimmer light and to a greater extent than the adaxial stomata. The abaxial and adaxial stomata of S. pennellii respond similarly to light incident on the adaxial epidermis and are less open at all quantum flux densities than comparable stomata of L. esculentum. The patterns of response to light of the abaxial and adaxial stomata of the chimera were practically identical to those of L. esculentum, and quite unlike those of S. pennellii. Thus, the pattern of stomatal light response in the chimera was regulated by the L. esculentum mesophyll. The reduction in stomatal frequency of the chimera, which was regulated by the epidermis of S. pennellii, contributed to the 40% difference in leaf diffusive resistance between the plants in moderate light.     

Short-term salt tolerance mechanisms in differentially salt tolerant tomato species

The physiological changes induced by a daily increase of NaCl level, over a period of 4 d, were studied in leaves of the salt-sensitive cultivated tomato species Lycopersicon esculentum and its wild salt-tolerant relative Lycopersicon pennellii. A higher solute contribution to the osmotic adjustment was observed in NaCl-treated leaves of L. pennellii than in those of L. esculentum. This response together with the higher accumulation of inorganic solutes in the wild species and of organic solutes in the cultivated species verified the different salt tolerance mechanisms operating in the two species in the short-term. With regard to the changes induced by salt stress on the free polyamine levels, the putrescine and spermine levels increased with salinity, whereas the spermine levels decreased in both tomato species; nevertheless, the main difference between the two species lays in an earlier and greater accumulation of putrescine induced by salinity in L. pennellii than in L. esculentum. The changes in putrescine levels were associated to changes in amino acids related to its synthesis, and the changes were different in both species. In L. esculentum, the high concentrations of some intermediate compounds (glutamate and arginine) were related to the low accumulation rate of both proline and putrescine. In contrast, in L. pennellii, important reductions in glutamate and arginine levels were found at the end of the salinization period. Moreover, in this last situation, a decline in the putrescine level ran parallel to a high proline accumulation, which suggests that the higher the stress level, the higher the deviation of glutamate to proline occurring in the salt tolerant species. It could be concluded that an early accumulation of the diamine putrescine seems to be associated with salt tolerance in the short-term.     

Genetic variability in Lycopersicon species and their genetic relationships

Summary Genetic diversity has to be described and measured in order to establish breeding strategies and manage genetic resources. It is also fundamental to develop a comparative intraspecific study before attempting to discuss and conclude any phylogenetic relationship. The genetic variability of Lycopersicon species was studied using starch gel electrophoresis of 11 enzymatic systems in a hierarchical fashion. The species with the greatest genetic variability are L. chilense, L. peruvianum and L. pennellii, mainly due to the within-line component. L. chmielewskii, L. parviflorum and L. pimpinellifolium show an intermediate total variability and their between-component clearly predominates over the within-component. The least variable species are L. cheesmanii and L. esculentum. Cluster analysis resulted in three main groups: one formed by the cultigen, L. pimpinellifolium, L. cheesmanii and L. peruvianum;another by two species with self-incompatibility systems, L. pennelli and L. chilense; and another by two autogamous species L. chmielewskii and L. parviflorum. With respect to L. esculentum the farthest related species is Solanum rickii and the closest, L. pimpinellifolium.     

Study of the three-genome hybridLycopersicon esculentum Mill. —L. chilense Dun. —L. peruvianum var ‘humifusum’ Mill. and its use as a source for resistance

L. peruvianum var humifusum is reproductively the most isolated of the species of the genusLycopersicon. It can be crossed with the cultivated tomato usingL. chilense as an intermediary. After a series of backcrosses of the three-genome hybrid F₁ (L. esculentum ×L. chilense) ×L. peruvianum var humifusum withL. esculentum, accompanied by selection for resistance to some economically important diseases, several lines were established. One of these lines, Cm 180, which showed resistance toClavibacter michiganensis subsp.michiganensis, was subjected to genetic analysis. This resistance was found to be controlled by a single dominant gene (Cm) that was not allelic to the gene originating fromL. hirsutum f.glabratum. ThisCm gene was genetically mapped on chromosome 4. The germ plasm ofL. peruvianum var humifusum in combination withL. chilense was transferred intoL. esculentum. Different breeding lines possessing resistance to various diseases and pests could be developed from this material.     

Inheritance and genetic mapping of resistance to Alternaria alternata f. sp. lycopersici in Lycopersicon pennellii

The fungal pathogen Alternaria alternata f. sp. lycopersici produces AAL-toxins that function as chemical determinants of the Alternaria stem canker disease in the tomato (Lycopersicon esculentum). In resistant cultivars, the disease is controlled by the Asc locus on chromosome 3. Our aim was to characterize novel sources of resistance to the fungus and of insensitivity to the host-selective AAL-toxins. To that end, the degree of sensitivity of wild tomato species to AAL-toxins was analyzed. Of all members of the genus Lycopersicon, only L. cheesmanii was revealed to be sensitive to AAL-toxins and susceptible to fungal infection. Besides moderately insensitive responses from some species, L. pennellii and L. peruvianum were shown to be highly insensitive to AAL-toxins as well as resistant to the pathogen. Genetic analyses showed that high insensitivity to AAL-toxins from L. pennellii is inherited in tomato as a single complete dominant locus. This is in contrast to the incomplete dominance of insensitivity to AAL-toxins of L. esculentum. Subsequent classical genetics, RFLP mapping and allelic testing indicated that high insensitivity to AAL-toxins from L. pennellii is conferred by a new allele of the Asc locus.     

Chromosome association and pollen fertility of parental and interspecific hybrids of Lycopersicon esculentum X L. hirsutum and L. pennellii

Cytological studies were carried out on two wild species (L. hirsutum and L. pennellii) and the cultivated species (L. esculentum) of tomato and their F1 hybrids. Both parents and hybrids show a diploid chromosome number of 2n=24. The meiotic behaviour of the cultivated species showed a high degree of chromosome homology resulting in a high level of chiasmata frequency per bivalent. In contrast, the two wild species showed a slight increase in uniyalent frequency and a decrease in bivalent formation and chiasmata frequency. The meiotic behaviour of the hybrids showed a high level of univalents and low levels of bivalents as well as trivalents. Highly significant decreases in chiasmata frequency and increases in meiotic abnormalities, especially in the L. esculentum X L. pennellii hybrid, also were detected. The high meiotic irregularity and low chiasmata frequency recorded in the second hybrid indicated the disharmony and difference between its parental genomes and also served to predict its sterility. With regard to degree of pairing recorded in the hybrids, there is a possibility that sterility in such cases may refer to genetic factors in addition to the previously mentioned reasons. Pollen fertility showed no great difference between L. esculentum and L. hirsutum and their F1 hybrid, but a significant decrease was recorded in the L. esculentum X L. pennellii hybrid, which was clearly associated with high meiotic irregularity, low chiasmata frequency and chromosome association.     

Trichome-borne and artificially applied acylsugars of wild tomato deter feeding and oviposition of the leafminer Liriomyza trifolii

Oviposition and adult feeding of the leafminer Liriomyza trifollii (Burgess) (Diptera, Agromyzidae) on Lycopersicon pennellii (Corr.) D'Arcy and its F₁ hybrid with Lycopersicon esculentum (Mill.) was significantly less than that on the cultivated tomato, L. esculentum. The resistance of L. pennellii and the F₁ was reduced following rinsing of foliage with ethanol. Resistant attributes of L. pennellii were transferred to L. esculentum through appression of L. pennellii foliage to L. esculentum leaflets. Application of purified 2,3,4-tri-O-acylglucoses (the principal component of type IV glandular trichome exudate of L. pennellii) to L. esculentum significantly decreased feeding and oviposition on L. esculentum leaflets by 61–99%. Therefore the principal mechanism of resistance to this leafminer by L. pennellii is the secretion of these acylglucoses. Dose response analysis of acylglucoses applied to L. esculentum shows that dosages as low as 10% those found on L. pennellii provide large reductions (91%) in leaf punctures and mines.     

Variations in the levels of major free cytokinins and free abscisic acid during tuber development of sweet potato

Changes in two plant growth substances were examined throughout tuber development of sweet potato (Ipomoea batatas Lam. cv. Minamiyutaka). Major free cytokinins [t-zeatin riboside, N⁶-(²-isopentenyl)adenosine and 6-(3-methyl-2-butenyl amino)purine glucoside] and free abscisic acid in tubers were determined by HPLC and GC-ECD. An increase int-ZR almost coincided with the onset of rapid tuberization 30–90 days after planting the vine cuttings. The levels of i⁶Ado and ABA were much lower than that oft-ZR throughout tuber development. The maximum level of i⁶Ado preceded the maximum oft-ZR. The level of iPG was higher than that oft-ZR, and the pattern of changes in the level was more complex. The maximum level of iPG reached about 270 g kg^–1 fresh weight. Varied changes in the low levels of ABA appeared not to be related to tuber development. Longitudinal distribution of the cytokinins and ABA in the developing tubers showed that levels oft-ZR were higher in parts of the proximal side of the stem than in other, lower parts of the tubers.     

Expression of unilateral incompatibility in pollen of Lycopersicon pennellii is determined by major loci on chromosomes 1, 6 and 10

Summary We have previously described gene introgression from the wild nightshade Solanum lycopersicoides into tomato (Lycopersicon esculentum) through the use of either diploid or sesquidiploid hybrids (the latter consisting of two genomes of L. esculentum and one genome of S. lycopersicoides). Both types of intergeneric hybrids display pollen sterility, but workable ovule fertility. Unilateral incompatibility prevents their direct hybridization with staminate L. esculentum. Pollen of a self-compattible form of the related wild species L. pennellii is compatible with pistils of L. esculentum x S. lycopersicoides hybrids. This trait was backcrossed from L. pennellii to L. esculentum in order to develop bridging lines that could be used to obtain progeny from the intergeneric hybrids and to study the inheritance of bridging ability. In progeny of L. esculentum x S. lycopersicoides hybrids pollinated with L. pennellii-derived bridging lines, preferential transmission of L. pennellii alleles was observed for certain isozyme and RFLP markers on chromosomes 1, 6 and 10. The skewed segregations suggest linkage to three major pollen-expressed compatibility loci. This was confirmed by observations of pollen tube growth, which indicated that compatibility with pistils of the diploid intergeneric hybrid occurred only in bridging lines at least heterozygous for the L. pennellii markers on chromosomes 1, 6 and 10. Compatibility with the sesquidiploid hybrid required only the chromosome 1 and 6 loci, indicating an apparent effect of gene dosage on expression of incompatibility in the pistil. In an F₂ L. esculentum x L. pennellii population, preferential transmission of L. pennellii alleles was observed for the same markers on chromosomes 1 and 10, as well as other markers on chromosomes 3, 11, and 12, but not 6. The chromosome 1 pollen compatibility locus maps to or near the S-locus, which determines S-allele specificity. The results are discussed in relation to existing genetic models for unilateral incompatibility, including the possible involvement of the S-locus.     

Feeding behavior of potato aphid affected by glandular trichomes of wild tomato

Mortality of the potato aphid, Macrosiphum euphorbiae (Thomas), on Lycopersicon pennellii (Corr.) D'Arcy and its F₁ hybrid with Lycopersicon esculentum Mill. was significantly greater than that on L. esculentum. Physical entrapment was not the sole mechanism of resistance in L. pennellii since few late instar aphids were found trapped in the sticky glandular exudate of the type IV trichomes; entrapment could, however, affect survival of early instars. Aphid settling on L. pennellii was dramatically less than that on L. esculentum, suggesting that starvation may have contributed to high mortality. Compared to L. esculentum, aphid feeding behavior on L. pennellii and the F₁ was characterized by a delay in the time to first probe, a reduction in the number of probes, and a decrease in the total proportion of time spent feeding. Removal of the glandular exudate of the type IV trichomes from L. pennellii resulted in a decrease in preprobe time and an increase in both the number of probes and the percent of time spent probing. Transfer of glandular trichome exudate of L. pennellii to leaflets of L. esculentum resulted in an increase in resistance as measured by these three parameters.

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Zusammenfassung Die Absterberate der Kartoffellaus, Macrosiphum euphorbiae Thomas, auf Lycopersicon pennellii (Corr.) D'Arcy, sowie auf der Kreuzung L. esculentum Mill. und L. pennellii, war deutlich grösser als auf L. esculentum. Das mechanische Verfangen der Läuse war nicht der Hauptgrund der Resistenz von L. pennellii. Wenige tote Läuse wurden in dem klebrigen Sekret der Typus IV Trichome gefunden. Auf L. pennellii siedelten sich die Läuse in viel geringerer Zahl an als auf L. esculentum. Dies führte zum Schluss, dass Verhungern eine der Ursachen der hohen Mortalität der Läuse war. Im Vergleich zum Saugverhalten auf L. esculentum war das Saugverhalten auf L. pennellii, wie auch auf F₁, durch Folgendes gekennzeichnet 1) Verspätung des ersten Stichversuchs, 2) Verminderung der Stichversuche pro Zeiteinheit und 3) Verminderung des Zeitanteils, der zum Saugen verwendet wurde. Die Entfernung des Sekrets der Typus IV Trichomen auf L. pennellii verursachte 1) eine kürzere Zeitspanne vor dem ersten Stichversuch, 2) eine Vergrösserung der Anzahl Stichversuche pro Zeiteinheit, 3) eine Verlängerung der Saugzeit. Die Uebertragung des Sekretes von L. pennellii auf Blätter von L. esculentum verbesserte deren Resistenz gegen Blattläuse gemessen mit den genannten drei Kriterien.