Population differentiation and the effects of herbivory and sand compaction on the subterranean growth of a desert lily

Differences in level of herbivory can select for local adaptation and genetic differentiation of plant populations in different environments. Mean bulb depth of the desert lily Pancratium sickenbergeri, differs considerably among populations differing in the level of herbivory by the dorcas gazelle. The gazelle digs in the sand to remove most of the bulb of the lily. Deeper bulbs have less material removed by herbivory than shallow bulbs and have higher fitness. A possible confounding factor is the degree of sand compaction, which may retard the downward growth of the bulb. We conducted a common garden experiment with 2 sand types with seeds from source populations with different levels of herbivory. There was a large genetic difference among populations. Two of 3 analyses indicated that there was an interaction between population and sand type, indicating that there is a heritable component of plasticity.     

Leaf compensatory growth as a tolerance strategy to resist herbivory in Pancratium sickenbergeri

We examined the effects of leaf herbivory by the dorcas gazelle, Gazella dorcas, on the compensatory growth of the geophyte Pancratium sickenbergeri (Amaryllidaceae) in the Negev desert, Israel. In three populations exposed to different levels of herbivory, we removed different amounts of photosynthetic leaf area from plants in five clipping treatments: 0, 25, 50%-dispersed over all leaves, 50%-entire area of half the leaves, 100%. The population with the lowest level of herbivory showed the lowest relative regrowth rate after clipping. In the population with a constantly high level of herbivory, plants in intermediate-clipping treatments overcompensated in leaf area after clipping. For all the populations, clipped plants produce more new leaves than unclipped plants. In the population with the highest level of herbivory, clipping treatments did not have a significant effect on the number of fruits per plant. In addition, we did not find a trade-off between investments in growth and reproduction in this population. Our results indicated that, in the desert lily, herbivores may select for plant mechanisms that compensate after damage as a tolerant strategy to maintain fitness.     

Patterns of <Emphasis Type="Italic">Azteca</Emphasis> ants’ defence of <Emphasis Type="Italic">Cecropia</Emphasis> trees in a tropical rainforest: support for optimal defence theory

Optimal defence theory (ODT) predicts that, whereas high risk of herbivory should select for high constitutive levels of defence, induced defences should be more advantageous in environments with a low probability of herbivory. In the present field study, conducted on the Azteca–Cecropia ant–plant system in a Neotropical rainforest, we evaluated whether the constitutive and induced ant defence of leaves are directly and inversely related to an estimate of herbivory risk, respectively. To assess the constitutive level of Azteca defence in Cecropia obtusifolia trees, we recorded the number of ants patrolling undamaged leaves. To evaluate the induced level of Azteca defence, the same leaves were subjected to simulated herbivory by punching circular holes in them. We recorded the maximum number of ants patrolling the damaged leaves from 2 to 15 min after damage. Past herbivory (% defoliation of old leaves) was assumed to indicate a risk of herbivory. Regression analyses showed that, whereas the constitutive level of ant patrolling was positively associated with the magnitude of herbivory on old leaves, there was a negative association between the magnitude of induced ant defence and past herbivory. These preliminary results lend support to ODT.     

Distinguishing signal from noise: Long-term studies of vegetation in Makhtesh Ramon erosion cirque,Negev desert,Israel

We consider three case studies of long-term plant population dynamics in Makhtesh Ramon erosion cirque in the central Negev desert of Israel. We show that rainfall is the major driving variable in this system, and that it creates large temporal and spatial variation in plant species diversity and vegetation community composition. This variability makes it extremely difficult to distinguish `signal' (= pattern in vegetation) from `noise' (random spatial and temporal variance). Our long-term vegetation studies in permanent plots arranged along the length of the cirque, initiated in 1990 and continuing, show that there is high spatial and temporal variance in plant species' incidences and abundances. This is particularly true of annual plant species. However, using pairs of fenced and unfenced plots arranged along the major abiotic gradient, altitude (which mirrors changes in rainfall), we were able to tease apart the effects of variance in rainfall and herbivory. We found significant negative effects of herbivory by the re-introduced Asiatic wild ass (Equus hemionus) on plant cover and on vegetation community composition. In a study of the effects of herbivory by dorcas gazelles (Gazella dorcas) on the lily Pancratium sickenbergeri, we found that there was little inter-annual fluctuation in lily population size in two sand dunes in Makhtesh Ramon in spite of the high percentage of lilies that is removed by the gazelles and the almost complete herbivory of flowers by these herbivores. This result indicates that the dune lily populations may be maintained by dispersal of seed from other lily populations elsewhere in Makhtesh Ramon where gazelles are rare or absent.     

The effects of herbivory and resource variability on the production of a second inflorescence by the desert lily, Pancratium sickenbergeri

We investigated the effects of herbivory by the dorcas gazelle, Gazella dorcas, on the production of a second inflorescence in a desert lily, Pancratium sickenbergeri (Amaryllidaceae), after the first inflorescence was eaten. In three populations exposed to different levels of herbivory, we conducted an inflorescence-clipping experiment in the flowering seasons of 1997, 1998 and 1999. The experiment was performed on inflorescences at both the emerging stage and the anthesis stage. Plants in each control group were undamaged. Clipped plants had a greater probability of producing a second inflorescence stalk than unclipped plants. In all the populations, the probability of producing a second inflorescence was greater when the first inflorescence was cut in the emerging stage than when it was cut in anthesis, indicating that there was a cost of compensation. In the population with the highest level of herbivory, the production of a second inflorescence stalk was related to resource availability in all 3 years. Plants produced as many flowers or fruits on the second stalk as on the first, indicating that potential reproductive output is double that usually achieved. Although we found no evidence for a trade-off between investment in first and second inflorescence stalks, the lower probability of producing a second inflorescence if heavy investment had been made in the first inflorescence indicates that compensation is costly. These results contradict the mutualism hypothesis that herbivory is beneficial to plants and provide support for the trade-off hypothesis that benefits of herbivory are proximal and are attained at an evolutionary cost.     

Evidence for rapid evolutionary change in an invasive plant in response to biological control

We present evidence that populations of an invasive plant species that have become re‐associated with a specialist herbivore in the exotic range through biological control have rapidly evolved increased antiherbivore defences compared to populations not exposed to biocontrol. We grew half‐sib families of the invasive plant Lythrum salicaria sourced from 17 populations near Ottawa, Canada, that differed in their history of exposure to a biocontrol agent, the specialist beetle Neogalerucella calmariensis. In a glasshouse experiment, we manipulated larval and adult herbivory to examine whether a population's history of biocontrol influenced plant defence and growth. Plants sourced from populations with a history of biocontrol suffered lower defoliation than naïve, previously unexposed populations, strongly suggesting they had evolved higher resistance. Plants from biocontrol‐exposed populations were also larger and produced more branches in response to herbivory, regrew faster even in the absence of herbivory and were better at compensating for the impacts of herbivory on growth (i.e. they exhibited increased tolerance). Furthermore, resistance and tolerance were positively correlated among genotypes with a history of biocontrol but not among naïve genotypes. Our findings suggest that biocontrol can rapidly select for increased defences in an invasive plant and may favour a mixed defence strategy of resistance and tolerance without an obvious cost to plant vigour. Although rarely studied, such evolutionary responses in the target species have important implications for the long‐term efficacy of biocontrol programmes.     

The Initiation and Growth of Tulip Bulbs

Tulip mother bulbs contain daughter bulbs, and second-generationdaughter bulbs are initiated within these daughter bulbs, fromFebruary to July. Initiation takes place progressively, thefirst-initiated bulbs being in the axils of the outermost scales.The innermost second-generation daughter bulbs are finally initiatedat the same time as the flower within the daughter bulb. Thelatest-formed bulbs grow much more rapidly than the others,and soon become larger than the first-formed ones. Later inthe season, growth of daughter bulbs falls into three phases,corresponding roughly with autumn, winter, and spring; growthis rapid in autumn and spring, and slower in winter. The ratesof daughter bulb growth in the autumn are affected by the amountof suppression by the mother bulb apex, but later these differencesdisappear. The durations of the phases of growth also vary withthe position of the daughter bulb. The outer daughter bulbsgrow for a longer time during the autumn phase than the innerones, so that differences in size due to variation in growthrate are partly nullified. In early spring, all daughter bulbsresume rapid growth simultaneously. Final bulb weight is thenapproached asymptotically, with the innermost bulbs having thehighest weights.     

Testing the optimal defence hypothesis for two indirect defences: extrafloral nectar and volatile organic compounds

Many plants respond to herbivory with an increased production of extrafloral nectar (EFN) and/or volatile organic compounds (VOCs) to attract predatory arthropods as an indirect defensive strategy. In this study, we tested whether these two indirect defences fit the optimal defence hypothesis (ODH), which predicts the within-plant allocation of anti-herbivore defences according to trade-offs between growth and defence. Using jasmonic acid-induced plants of Phaseolus lunatus and Ricinus communis, we tested whether the within-plant distribution pattern of these two indirect defences reflects the fitness value of the respective plant parts. Furthermore, we quantified photosynthetic rates and followed the within-plant transport of assimilates with (13)C labelling experiments. EFN secretion and VOC emission were highest in younger leaves. Moreover, the photosynthetic rate increased with leaf age, and pulse-labelling experiments suggested transport of carbon to younger leaves. Our results demonstrate that the ODH can explain the within-plant allocation pattern of both indirect defences studied.     

Cascading effects of induced terrestrial plant defences on aquatic and terrestrial ecosystem function

Herbivores induce plants to undergo diverse processes that minimize costs to the plant, such as producing defences to deter herbivory or reallocating limited resources to inaccessible portions of the plant. Yet most plant tissue is consumed by decomposers, not herbivores, and these defensive processes aimed to deter herbivores may alter plant tissue even after detachment from the plant. All consumers value nutrients, but plants also require these nutrients for primary functions and defensive processes. We experimentally simulated herbivory with and without nutrient additions on red alder (Alnus rubra), which supplies the majority of leaf litter for many rivers in western North America. Simulated herbivory induced a defence response with cascading effects: terrestrial herbivores and aquatic decomposers fed less on leaves from stressed trees. This effect was context dependent: leaves from fertilized-only trees decomposed most rapidly while leaves from fertilized trees receiving the herbivory treatment decomposed least, suggesting plants funnelled a nutritionally valuable resource into enhanced defence. One component of the defence response was a decrease in leaf nitrogen leading to elevated carbon : nitrogen. Aquatic decomposers prefer leaves naturally low in C : N and this altered nutrient profile largely explains the lower rate of aquatic decomposition. Furthermore, terrestrial soil decomposers were unaffected by either treatment but did show a preference for local and nitrogen-rich leaves. Our study illustrates the ecological implications of terrestrial herbivory and these findings demonstrate that the effects of selection caused by terrestrial herbivory in one ecosystem can indirectly shape the structure of other ecosystems through ecological fluxes across boundaries.     

Latitudinal patterns of herbivore pressure in a temperate herb support the biotic interactions hypothesis

The longstanding biotic interactions hypothesis predicts that herbivore pressure declines with latitude, but the evidence is mixed. To address gaps in previous studies, we measured herbivory and defence in the same system, quantified defence with bioassays, and considered effects of leaf age. We quantified herbivory and defence of young and mature leaves along a continental gradient in eastern North America in the native herb Phytolacca americana L. Herbivory in the field declined with latitude and was strongly correlated with lepidopteran abundance. Laboratory bioassays revealed that leaf palatability was positively correlated with latitude of origin. Young leaves were more damaged than mature leaves at lower latitudes in the field, but less palatable in bioassays. Both defence and palatability displayed non‐linear latitudinal patterns, suggesting potential mechanisms based on biological or climatic thresholds. In sum, observational and experimental studies find patterns consistent with high herbivore pressure and stronger plant defences at lower latitudes.     

BULB ORGANIZATION IN ALLIUM: SOME SPECIES OF THE SECTION MOLIUM

Mann , Louis K. (U. California, Davis.) Bulb organization in Allium: some species of the section Molium. Amer. Jour. Bot. 47(9) : 765–771. Illus. 1960.—Bulb development and organization are described for Allium neapolitanum, A. erdelii, A. hirsutum, A. subhirsutum, A roseum, and A. zebdanense, species of Molium, a Mediterranean section of the genus. These species begin growth in the late fall or midwinter, flower in March and April, and are dormant throughout the summer. The plant axis is commonly a sympodium made up of one or more branches terminated by inflorescences and a final branch ending in a large renewal bulb.. This bulb and one to many increase bulbs are formed each season. The bulbs mature by flowering time or shortly thereafter and the other parts—stem, leaves, roots, etc.—of the current season's plant die. Little change is evident within the bulbs during the summer rest period, but in September and October (or later in one species) the inflorescence and renewal bulb primordia appear. Five distinct types of leaves make up the bulb, including foliage leaves, food-storage leaves, and leaves which develop a heavy sclerotic layer around the bulb. The similar bulb structure of the 6 species indicates their close relationship. The 2 leaves which form the protective layer around the bulb become fused by the time they reach maturity, a condition apparently unique for the section Molium.     

Herbivory limits the yellow water lily in an overgrown lake and in flowing water

Aquatic macrophytes with floating leaves are often key ecological species that affect entire aquatic ecosystems. Here we describe an investigation of the importance of insect herbivory for population growth and leaf senescence in the yellow water lily (Nuphar lutea). In order to gain a general picture of the importance of herbivory under different conditions, we experimentally manipulated herbivory in a large lily population in natural still water and observed the natural development of 32 smaller populations in flowing water. Herbivory drastically increased leaf senescence, reducing leaf density. In the still water, over one summer, leaf density increased by a factor of 1.23 in the presence of water lily leaf beetles and 1.61 when herbivory was eliminated. In flowing water, population growth was restricted mainly by leaf crowdedness, which limited large dense populations. Herbivory by water lily leaf beetles also had a limiting effect on yellow water lily, again mainly in large dense populations. Small populations supported a lower density of beetles. Previous studies have not addressed population-level responses of vascular plants with floating leaves. Our results suggest that herbivory can result in greater light penetration into the water and reduce “enemy-free space” for aquatic species that find such space in water lily stands. We suggest that the water lily leaf beetle should be considered an “ecological engineer.”     

Flowering pathway is regulated by bulb size in Lilium longiflorum (Easter lily)

Lilium longiflorum (Easter lily) vegetative propagation occurs through production of underground bulbs containing apical and axillary meristems. In addition, sexual reproduction is achieved by flowering of elongated shoots above the bulb. It is generally accepted that L. longiflorum has an obligatory requirement for vernalisation and that long day (LD) regime hastens flowering. However, the effect of bulb size and origin, with respect to axillary or apical meristems on flowering, as well as the interactions between these meristems are largely unknown. The aim of this study was to explore the effect of bulb size, vernalisation and photoperiod on L. longiflorum flowering. To this end, we applied vernalisation and photoperiod treatments to the different bulb sizes and used a system of constant ambient temperature of 25 °C, above vernalisation spectrum, to avoid cold‐dependent floral induction during plant growth. Vernalisation and LD hasten flowering in all bulbs. Large, non‐vernalised bulbs invariably remained at a vegetative stage. However, small non‐vernalised bulbs flowered under LD conditions. These results demonstrate for the first time that cold exposure is not an obligatory prerequisite for L. longiflorum flowering, and that an alternative flowering pathway can bypass vernalisation in small bulbs. We suggest that apical dominance interactions determine the distinct flowering pathways of the apical and axillary meristems. Similar floral induction is achieved in propagated bulblets from scaling. These innovative findings in the field of geophyte floral induction represent valuable applicative knowledge for lily production.     

Trade-offs in non-native plant herbivore defences enhance performance

Non-native plants are typically released from specialist enemies but continue to be attacked by generalists, albeit at lower intensities. This reduced herbivory may lead to less investment in constitutive defences and greater investment in induced defences, potentially reducing defence costs. We compared herbivory on 27 non-native and 59 native species in the field and conducted bioassays and chemical analyses on 12 pairs of non-native and native congeners. Non-natives suffered less damage and had weaker constitutive defences, but stronger induced defences than natives. For non-natives, the strength of constitutive defences was correlated with the intensity of herbivory experienced, whereas induced defences showed the reverse. Investment in induced defences correlated positively with growth, suggesting a novel mechanism for the evolution of increased competitive ability. To our knowledge, these are the first linkages reported among trade-offs in plant defences related to the intensity of herbivory, allocation to constitutive versus induced defences, and growth.     

Potential defence from herbivory by ‘dazzle effects’ and ‘trickery coloration’ of leaf variegation

The very conspicuous dazzle coloration invented for naval defence during World War I was used in pre‐radar days to mislead attackers of naval units about vessel size, type, speed, and direction. Among several potential types of defences, it is proposed that zebra‐like white leaf variegation may defend leaves and other plant organs from herbivory as a result of dazzle effects. Two different dazzle effects may be involved in defending plants from herbivory, making it hard for herbivores (1) to decide where, in a three‐dimensional space, to bite the leaves (large herbivores) and (2) to land on them (insects). In addition, the related types of leaf coloration described in the present study, comprising parallels of military defensive trickery naval painting, may also deceive herbivores about the actual shape, location, and identity of leaves. Some of these visual defences may operate at the same time as other visual defences, such as aposematism, or serve various physiological functions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 692–697.     

Effects of exogenous gibberellins and paclobutrazol on floral stalk growth of tulip sprouts isolated from cooled and non-cooled tulip bulbs

The involvement of gibberellins (GAs) in the regulation of floral stalk elongation and flower development has been studied in tulip. The biological activity of GA₄ and GA₉, both endogenous in tulip bulb sprouts, and GA₁, was tested in vitro on sprouts of cooled and non-cooled tulip bulbs ( Tulipa gesneriana L. cv. Apeldoorn), in the presence or absence of the GA biosynthesis inhibitor paclobutrazol. At early starting dates of incubation, floral stalks from both cooled and non-cooled bulbs hardly showed any elongation in the absence of exogenous GA. Paclobutrazol had no effect on floral stalk elongation, and the response to GAs of sprouts from cooled bulbs was greater than that of sprouts from non-cooled bulbs. At later starts of incubation, considerable floral stalk elongation occurred without GA application. Paclobutrazol inhibited this floral stalk elongation, and the growth of sprouts from both cooled and non-cooled bulbs was stimulated by GA application. The effect of paclobutrazol was reversed by simultaneous application of GA₄ or GA₉. Application of GA with and without paclobutrazol resulted in the same elongation of the floral stalk, indicating the absence of substantial side effects of the inhibitor. The isolated sprouts did not develop a full-grown flower without the addition of GA. GA₄ was more effective than GA₉ in stimulating this flower development. The results demonstrate that both sprouts from cooled and non-cooled bulbs are responsive to exogenous GAs in vitro, and may be a site of GA biosynthesis.     

Dietary Overlap of Przewalski's Gazelle,Tibetan Gazelle,and Tibetan Sheep on the Qinghai-Tibet Plateau

Abstract: We studied food composition and overlap among sympatric Przewalski's gazelle (Procapra przewalskii), Tibetan gazelle (P. picticaudata), and Tibetan domestic sheep (Ovis aries) in the Upper Buha River, Qinghai-Tibet Plateau, using microscopic fecal analysis. The 2 gazelles forage mainly on plants from Leguminosae and Compositae families and domestic sheep feed mainly on Gramineae and Cyperaceae. The dietary overlap index between the 2 gazelles and Tibetan domestic sheep increased from 0.43 to 0.58 during the plant-growing period to 0.76–0.77 during the plant-withering period, which indicated competition for foods intensified during the food-limited season. Although the 2 gazelle ate similar diets, they might avoid food competition by occupying different foraging areas. We suggest reducing numbers of overwintering Tibetan domestic sheep on pastures to lessen survival pressure on the endangered gazelles during winters.     

Red leaf margins indicate increased polygodial content and function as visual signals to reduce herbivory in Pseudowintera colorata

Red-pigmented leaf margins are common, but their functional significance is unknown. We hypothesized that red leaf margins reduce leaf herbivory by signalling to herbivorous insects the presence of increased chemical defences. Leaves were collected from a natural population of Pseudowintera colorata. Margin size, herbivory damage, anthocyanin content and concentrations of polygodial, a sesquiterpene dialdehyde with antifeedant properties, were quantified. Feeding trials involving larvae of Ctenopseustis obliquana, a generalist herbivore, were conducted on red- and green-margined P. colorata leaves in darkness, or under white, green or red light. Leaves with wider red margins contained higher concentrations of polygodial and anthocyanins, and incurred less natural herbivory. In trials under white light, C. obliquana consumed disproportionately more green- than red-margined leaf laminae. Larvae exhibited no feeding preference when light was manipulated such that leaf colour discrimination was impaired. Red leaf margins provide a reliable and effective visual signal of chemical defence in P. colorata. Ctenopseustis obliquana larvae perceive and respond to the colour of the leaf margins, rather than to olfactory signals. Our study provides direct experimental evidence for aposematic coloration in red leaves.     

Biological Control of the Bulb Mite, Rhizoglyphus robini , by the Predatory Mite, Hypoaspis aculeifer , on Lilies: Predator-Prey Dynamics in the Soil, under Greenhouse and Field Conditions

We tested the capacity of the soil-dwelling predatory mite, Hypoapsis aculeifer , to control mites attacking lily bulbs. Experiments in the greenhouse and in the field showed that in the absence of predatory mites populations of the bulb mite, Rhizoglyphus robini , on lily bulbs increased, whereas the release of predatory mites either slowed down the increase - as observed in the field - or caused the bulb mites populations to decrease - as observed in the greenhouse. In all cases the population of predatory mites increased as long as bulb mite densities were not too low. However, within the first week after predator release there was usually a sharp decline to 10-40% of the original number released. Greenhouse experiments on intact lily bulbs in pots, boxes and 1 m 2 plots with peat soil showed that when released in a ratio of 1 predator to 2 or 5 prey, the predatory mite, Hypoaspis aculeifer , suppressed populations of bulb mites to less than 10 individual per bulb within 6 weeks. Elimination of bulb mites was observed only when the predator-to-prey ratio at release was equal to 3:1. Field experiments in 2 m 2 plots with intact bulbs in rather compact sandy soil showed that when released in ratio of 1 predator to 1 or 2 prey, the predatory mite, H. aculeifer , did not cause the population of bulb mites to decrease, but it did reduce their population growth. The initial predator-to-prey ratios required to achieve suppression (ca 1:2) or elimination (3:1) in the soil environment are much higher than those required for bulb mite elimination when lily bulb scales were embedded in a medium of vermiculite (ca 1:20). Among the possible causes are: (1) the initial losses of predators in the greenhouse and even more so in the field due to mortality and/or emigration from the experimental plots; (2) the lower temperatures in the greenhouse and especially in the field, which slow down the growth and predation processes and thereby delay prey extinction; and (3) the spatial complexity of the soil environment which creates refuges for the bulb mites.