常绿阔叶林林下6种木本植物叶片非结构性碳水化合物的动态特征

植物叶片非结构性碳水化合物(NSC)不仅为植物代谢提供重要能量,还能一定程度上反映植物对外界环境的适应策略.以亚热带常绿阔叶林林下6种植物:茶(Camellia sinesis)、细枝柃(Eurya loquaiana)、润楠(Machilus pingii)、短刺米储(Castanopsis carlesii)、大叶山矾(Symplocos grandis)和薄叶山矾(Symplocos anomala)为对象,研究各物种抽枝展叶进程中叶片大小、比叶重(LMA)、光合色素以非结构性碳水化合物及其组分含量的动态变化,分析NSC组分之间及与光合色素与LMA间的相互关系,探讨展叶过程中引起叶片NSC差异的原因.结果表明:(1)各物种单叶面积随叶片展开而增加直至8月下旬达到最大,为9.20 (茶)-40.81 cm2 (大叶山矾);而LMA在展叶初期下降后随叶片展开逐步升高,直到次年1月下旬还在持续缓慢增加,因各物种不同,最大值为82.90-152.10 g/m2;光合色素则在展叶进程中逐渐增加,6月下旬达到较高值后在整个夏秋季维持较高的含量,次年1月有所下降.(2)在整个当年生叶片生长进程中,6种植物叶片可溶性糖含量总体上由展叶初期逐渐增加,而淀粉含量随着叶片的生长成熟逐渐降低;各物种NSC含量为87.00 (薄叶山矾)-163.35 mg/g(细枝柃),除大叶山矾和薄叶山矾外,NSC含量随着叶片生长进程逐渐增加;(3)各物种可溶性糖含量随着淀粉含量的增加而降低,叶绿素含量随着LMA的增加而显著增加,可溶性糖与LMA具有显著的正相关关系,而淀粉与LMA呈显著负相关(R~2=0.51-0.86,P0.004),表明展叶后期部分淀粉转化为可溶性糖,这与展叶后期林内光资源的可利用性以及植物的生理活动相关.综上所述,随着展叶进程,非结构性碳水化合物及其组分具有不同的变化规律,一方面与叶片增大增厚进程中的生理活动有关,另一方面也反映了叶片功能属性间的权衡关系,研究结果可为阐明亚热带常绿阔叶林林下木本植物展叶期的碳代谢提供理论基础,亦丰富了森林植物生活史对策理论.(图6参53)     

哀牢山12种木质藤本植物在不同生境下叶片解剖结构的比较

为了探讨藤本植物对不同生境的适应性,以哀牢山亚热带常绿阔叶林中12种常见的木质藤本植物为研究对象,包括6种林窗更新的、喜光或适应高光强生境的常绿蔷薇(Rosa longicuspis)、山羊桃(Actinidia callosa)、南蛇藤(Celastrus angulatus)、五风藤(Holboellia latifolia)、云南清风藤(Sabia yunnanensis)、圆锥悬钩子(Rubus paniculatus)和6种林缘或林内更新的、喜荫或适应低光强生境的冷饭团(Kadsura coccinea)、石宝茶藤(Euonymus vagans)、川西尾叶素馨(Jasminum urophyllum)、冠盖绣球(Hydrangea anomala)、肖菝葜(Heterosmilax japonica)和葡萄酸藤子(Embelia procumbens),使用石蜡切片法对其在林窗(全光照)、林缘(遮荫)和林内(荫生)3种生境下的植物叶片进行了解剖观察和比较分析,研究不同生境对这12种木质藤本植物叶片结构的影响。结果表明,(1)从林窗到林缘再到林内,12种木质藤本植物的叶片平均厚度、上表皮平均厚度、栅栏组织平均厚度、栅栏组织细胞平均长度和栅海比均呈现逐渐降低的趋势,且林窗更新的木质藤本植物的下降趋势大于林缘、林内更新的。(2)栅栏组织细胞层数及其长度随生境光强减弱显著减少和变短,导致栅栏组织平均厚度显著变小,可作为藤本植物响应生境变化的重要指标。(3)12种木质藤本植物的总体表型可塑性指数平均值为0.47,其中,6种林窗更新的藤本植物的总体表型可塑性指数平均值(0.58)大于6种林缘、林内更新的藤本植物的(0.36)。结果说明,木质藤本植物叶片解剖结构表现出对生境光强具有很强的适应性;同时,喜光或适应高光强的藤本植物对异质光生境的适应能力强于喜荫或适应低光强的藤本植物,这可能是前者能够在常绿阔叶林3种生境中具有更高分布频率,而后者多分布于林缘和林内生境的原因之一。     

林丹烟剂对毛竹林冠与林下层节肢动物共有种及其多样性的影响

共有种反映毗邻生境节肢动物群落的相互作用程度及其潜在互作途径的多样性.通过对施用林丹烟剂后毛竹林冠层与林下层节肢动物的系统定位调查,分析和比较了林丹烟剂干扰下两个林层的节肢动物共有种及其多样性.结果表明,以未防治和白僵菌防治为对照,施用林丹烟剂显著减少了林冠和林下层的共有种及其个体数,降低了两个林层中共有种的多样性;在物种方面主要表现为蜘蛛目、膜翅目、鞘翅目、双翅目和鳞翅目共有种减少;在个体方面,林下层主要表现为蜘蛛目、膜翅目、鞘翅目和双翅目共有种个体减少,林冠层主要表现为蜘蛛目、鞘翅目和双翅目共有种个体减少.但无论在物种还是个体方面,林丹烟剂对捕食性共有种的影响均最大.进一步的灰色关联分析表明,引起共有种多样性下降的主要原因在于共有物种的减少.因此认为,林丹烟剂干扰削弱了林下和林冠层节肢动物之间的相互作用,不利于林下生境促进林冠层生物多样性和增强群落对叶部害虫自然控制作用等生态功能的发挥.图1表6参22     

林窗大小对川西马尾松人工林林下物种多样性和生物量的影响

林窗是植被演替和森林循环的重要驱动力,研究林窗大小对物种多样性和生物量的影响有利于揭示林窗干扰在生态系统恢复中的作用.以川西莲花山马尾松人工林为研究对象,划分出5种不同面积(81-625 m~2)的林窗共计15个和3个非林窗(林下)作对照(CK),对其群落物种组成、物种多样性水平(丰富度指数D、Simpson优势度指数H’、Shannon-Wiener指数H、Pielou均匀度指数J_(sw))和林下生物量(灌木层生物量、草本层生物量)进行调查研究.结果表明:(1)在总面积为5 493 m~2的样地中,共记录到维管植物99种,隶属56科84属;草本层和灌木层物种数随林窗面积增加均呈单峰格局,前者的物种数变化较后者明显.(2)总的来看,林窗下的D、H、J_(sw)值较林下大,H’值较林下小,且400-450 m~2面积的林窗下灌木、草本各物种多样性指数值达到一个较高水平.(3)林窗内的生物量显著大于林下生物量,200-250 m~2林窗下的生物量最大.(4)群落各层次物种多样性指数与对应层次地上、地下、全株生物量的相关性均不显著.因此认为,大面积林窗(400-450 m~2)有利于增加马尾松人工林林下群落物种丰富度,提升林下群落物种多样性水平;而中等面积林窗(200-250 m~2)则有利于增加林下生物量.     

川西周公山柳杉人工林林窗大小对土壤理化性质和物种多样性的影响

以川西周公山柳杉人工林林中5种不同大小(50~100、100~150、200~250、400~450、600~650 m2)的林窗为研究对象,通过对林窗内土壤pH值、有机质含量等指标和物种多样性的研究,探讨天然林窗的形成对土壤各理化指标以及物种多样性的影响。结果表明:(1)随着林窗面积的增大,群落各层次的物种数呈现出先升高后降低的趋势,灌木层、草本层的Shannon-Wiener多样性指数(H)、Simpson优势度指数(H')和Pielou均匀度指数(Jsw)均表现为林窗林下;(2)0~20 cm土壤速效钾含量表现为林窗林下;(3)各物种多样性指数变化与土壤理化指标存在一定相关性,草本层物种丰富度指数、Shannon-Wiener多样性指数及Simpson优势度指数与土壤pH值呈显著相关关系(P0.05),物种丰富度指数、Shannon-Wiener多样性指数与速效钾含量呈显著相关关系(P0.05)。     

林窗对马尾松人工林植物多样性的影响

为了解人工抚育调控林窗对马尾松人工林植物多样性的影响,以长江上游低山丘陵区39年生的马尾松人工林为研究对象,人为砍伐形成不同大小(100-1 600 m2)的林窗共计21个,研究了林窗、林窗边缘、林下的植物多样性.结果表明:马尾松人工林林窗内外的丰富度指数R0(物种数)、Simpson生态优势度指数(SP)、Shannon-Wiener物种多样性指数(SW)差异显著(P0.05),其丰富度指数的大小顺序是林窗林窗边缘林下.不同大小林窗之间的丰富度指数也有显著差异(P0.05),但Simpson生态优势度指数、Shannon-Wiener物种多样性指数和Pielou均匀度指数(Jsw)差异均不显著.不同大小林窗边缘的丰富度指数、优势度指数、多样性指数和均匀度指数均没有显著差异(P0.05).综合来看,林窗的形成显著提高了马尾松人工林林下植物的丰富度,林窗大小对林下植物的物种组成和丰富度也有显著影响.     

林窗大小对2种针叶林更新效果的初步分析

为了解采伐林窗对2种人工林更新效果的影响,在浙江省丽水市白云山选择生境条件基本一致的马尾松林和杉木林,以林下为对照,比较了通过采伐获得的不同大小林窗(50～500 m~2)内人工和天然更新植物效果。结果表明:(1)10个人工更新树种在杉木林中生长优于马尾松林,马尾松林中枫香、无患子、山乌桕和细柄蕈树生长情况好于其他树种,杉木林中枫香、山乌桕、南方红豆杉、无患子和细柄蕈树生长情况好于其他树种。(2)林分类型显著影响南方红豆杉生长,特别是对树高的影响极为显著(P0.01),林窗大小对木荷地径和树高均有显著影响(P0.05),林分和林窗的交互作用对除朴树、闽楠和浙江楠外的其他树种均有显著影响。(3)不同大小林窗下天然更新植物物种数、Simpson生态优势度、Shannon多样性指数存在显著差异(P0.05),林分类型对物种数和Shannon多样性指数具有显著影响(P0.05),但对Simpson生态优势度没有影响。2种林分林窗间天然更新植物相似度介于0.4～0.9之间,同一林分林窗间物种相似度均大于林窗与对照之间。马尾松林阔叶化改造可采伐中等林窗(250 m~2),目标更新树种可选枫香、无患子、细柄蕈树和木荷;杉木林以大(500 m~2)或小林窗(50 m~2)为主,可选树种有枫香、南方红豆杉、山乌桕、无患子和细柄蕈树。     

林窗对川西亚高山6种植物凋落叶纤维素降解的影响

凋落物纤维素降解是森林生态系统碳循环的关键过程,其可能受到林窗直接或间接的影响.采用凋落物分解袋法,通过3年连续监测,研究川西亚高山6种常见木本植物凋落叶(岷江冷杉、红桦、四川红杉、方枝柏、康定柳和高山杜鹃)在不同林冠环境下(林窗中心、林缘林窗、扩展林窗和郁闭林下)冬季和生长季纤维素降解特征.结果显示:川西亚高山6种凋落叶分解过程中纤维素含量和残留率均呈现下降趋势,分解3年后纤维素含量为5.5%-12.1%且残留率下降至23.1%-44.6%.林窗中心处整体上具有较低的凋落叶纤维素含量和残留率,但这种差异性在分解3年后消失.相关性分析表明凋落叶纤维素含量和残留率与均温、初始纤维素、木质素含量和C/N值呈显著负相关,与冻融循环和初始有机溶性组分含量呈显著正相关.表明林窗的形成通过提供较高的温度或减少冻融循环的发生促进凋落叶纤维素的降解,但这种差异随着分解进程逐渐减弱,且受到凋落叶初始质量所代表的物种差异的影响.本研究表明林窗通过改变林内微环境促进凋落叶纤维素早期的降解,上述结果有助于进一步认识林窗在亚高山森林生态系统物质循环中的生态作用和意义.(图4表2参39)     

川西高山森林生态系统林下生物量及其随林窗的变化特征

作为森林生态系统的重要组成部分,林下植被及其残体的分布受到林冠层的影响,但迄今有关林窗对林下植被和残体生物量的影响尚无研究报道.于2013 年8 月2 日至20 日,以海拔3 600 m 的川西岷江冷杉原始林林下植被为研究对象,根据区域内的坡向和林分组成等因素设置3 个100 m×100 m 的典型样地,调查其生物量及其随林窗的变化特征.在每个样地内选择3 个大林窗,在林窗、林缘和林下分别设置3 个20 m×20 m 的样方,调查粗木质残体长度或高度、大小头直径、枯立木记录胸径、腐烂等级等;在林窗、林缘和林下分别设置3 个5 m×5 m 的样方,采用“收获法”收集样方内直径在2.5-10 cm之间的细木质残体和灌木生物量;在林窗、林缘和林下分别设置3 个1 m×1 m 的样方来调查凋落物储量和草本生物量;在1m×1 m 的样方内随机选择1 个20 cm×20 cm 的小样方来调查地被植物生物量.结果表明,（1）川西高山森林生态系统总生物量为72.75 t·hm^-2,其中林下生物量为67.92 t·hm^-2,占生态系统生物量的95.17%.活体植被以灌木为主,其生物量为9.81t·hm^-2;残体部分以粗木质残体为主,其储量为53.00 t·hm^-2;（2）林窗对灌木、草本、地被植物的影响各不相同,且不同物种的灌木生物量表现出不同的分布规律;草本生物量表现出明显的“边缘效应”,在林缘显著高于林下;林窗和林缘的地被植物生物量相对较低;（3）粗木质残体储量从林下到林窗呈现减小的趋势,但总体储量仍然较大,林窗和林缘的细木质残体储量高于林下.这些结果为认识高山森林生态系统林下生物量及其格局,以及林窗在森林生态系统的重要作用提供了基础理论依据.     

林窗边缘效应对马尾松和香樟凋落叶分解的影响

为了解马尾松(Pinus massoniana)人工林不同面积林窗边缘凋落叶分解的养分释放规律,采用凋落袋分解实验,以长江上游低山丘陵区人为采伐形成的马尾松人工林7种不同大小林窗(G1:100 m~2;G2:225 m~2;G3:400 m~2;G4:625 m~2;G5:900 m~2;G6:1 225 m~2;G7:1 600 m~2)为研究对象,以林下为对照,探讨马尾松和香樟(Cinnamomum camphora)凋落叶的质量损失率和养分释放率.结果表明:除钾释放率外,林窗边缘两种凋落叶的质量损失率和养分释放率均显著高于林下.在1.5年分解过程中,两种凋落叶的质量损失率和碳释放率呈持续上升趋势.分解前90 d,两种凋落叶的质量损失速率和养分释放速率最快;分解90-360 d后,部分林窗出现氮和磷的富集现象;180-270 d和1.5年后,部分林窗出现钾的富集现象.随林窗面积增大,除钾释放率外,两种凋落叶在中型林窗边缘(400、600和900 m~2)具有较高的质量损失率和养分释放率.在凋落叶分解过程中,质量损失率和碳、氮释放率与土壤含水率均呈显著正相关.中型林窗(400-900 m~2)对凋落物的分解具有更显著的边缘效应,而这种边缘效应与物种有一定关联.因此,在马尾松低效林改造过程中,可利用中型林窗(400-900 m~2)边缘这一显著作用,引入乡土阔叶树种加速马尾松人工林的物质循环,维持林分地力.     

Jasmonic acid treatment and mammalian herbivory differentially affect chemical defenses and growth of wild mustard (Brassica kaber)

Summary. Jasmonic acid (JA) is a wound-related hormone found in most plants that, when applied exogenously, can induce increases in levels of chemical defenses in patterns similar to those induced by mechanical damage or insect feeding. Relative to responses to insect and pathogen attack, chemical responses of herbaceous plants to mammalian herbivore attack have been little studied. In a field experiment, we compared the effects of JA treatment and naturally occurring mammalian herbivory on the expression of trypsin inhibitors, glucosinolates, peroxidase activity and growth of wild mustard (Brassica kaber). Exogenous JA significantly increased trypsin inhibitor activity and glucosinolate concentration, and moderately increased peroxidase activity in the eighth true leaves of five-week-old plants, relative to untreated controls. In contrast, levels of these chemical defenses in the eighth true leaves or in regrowth foliage of plants that had ∼80% of their leaf area removed by groundhogs (Marmota monax) did not differ from that in undamaged and untreated controls. Although exogenous JA significantly elevated levels of chemical defenses, it did not affect height of plants through the season and only slightly reduced time to first flower. Groundhog herbivory significantly reduced height and delayed or abolished flowering, but these effects were not substantial unless coupled with apical meristem removal. We hypothesize that the lack of effect of groundhog herbivory on chemical defenses may be due in part to the speed and pattern of leaf area removal by groundhogs, or physiological constraints caused by leaf area loss. Despite having no effect on chemical defense production, leaf area loss by groundhogs was more costly to growth and fitness than the effects of JA application in this study, but only substantially so if coupled with apical meristem removal. We suggest that in general, costs of defense production in plants are likely to be minimal when compared to the risk of losing large amounts of leaf area or primary meristematic tissue. Thus, if they are effective at deterring herbivory, the benefits of inducible defense production likely outweigh the costs in most cases. Received 20 December 2000; accepted 3 May 2001     

Assessing the antipredatory defensive strategies of Caribbean non-scleractinian zoantharians (Cnidaria): is the sting the only thing?

The relative importance of chemical, nematocyst, and nutritional defenses was examined for 18 species of Caribbean sea anemones (actinarians), zoanthids, and mushroom polyps (corallimorpharians) from the Florida Keys and the Bahamas Islands, 2008–2010. Feeding assays were performed using the fish Thalassoma bifasciatum with artificial foods containing crude organic extracts of cnidarian tissues. A novel behavioral assay using brine shrimp nauplii was used to assess nematocyst defenses. The nutritional quality of cnidarian tissues was examined using bomb calorimetry and soluble protein assays. In general, actinarians invested in nematocyst defenses, zoanthids in either nematocyst or chemical defenses, and corallimorpharians lacked both, except for 1 of 3 species that was chemically defended. Relative to other coral reef invertebrates, cnidarian tissues had similar caloric values but lower soluble protein concentrations. Trade-offs between chemical and nematocyst defenses were observed for 65% of species, while habitat and behavior provided a likely explanation for undefended species.     

A review of chemical ecology in poison frogs

Herein we review what is known about the chemical ecology of poison frogs with a focus on dendrobatid poison frogs. While five anuran families are known to have an alkaloid-derived chemical defense, the dendrobatids have been studied in greatest detail and provides chemical ecologists with a complex model system for understanding how chemical defenses operate in real time and may have evolved through evolutionary time. We describe the diversity of alkaloid defenses known from frogs, alkaloid sequestration, biosynthesis and modification, and we review what is known concerning arthropod sources for alkaloids. There is variation in nearly every attribute of the system and we try to describe some of the challenges associated with unraveling the complexities of this model system.     

Within-plant allocation of a chemical defense in Secale cereale. Is concentration the appropriate currency of allocation?

Summary. The among-leaves allocation of DIBOA, a hydroxamic acid associated with plant resistance, in the shoot of rye (Secale cereale) was evaluated over the vegetative development of the plant. The appropriateness of using the concentration of secondary metabolites, DIBOA in this case, as the parameter to evaluate defense allocation in plants is discussed. Both biological and statistical arguments are put forward to suggest that allocation of chemical defenses should refer to absolute content and not to concentration. Results showed that leaf age was significantly linked to leaf concentration of DIBOA, young leaves having higher concentrations. In contrast, leaf content of DIBOA, our proposed currency of allocation, was not significantly higher in younger leaves. Furthermore, a regression analysis showed that the DIBOA content of leaves was better explained by the leaf relative biomass (proportion of shoot biomass) than by leaf biomass itself. It is suggested that, rather than leaf age, leaf relative biomass is the major factor determining DIBOA allocation in rye shoots. It is proposed that studies addressing within-plant defense allocation should use chemical defense content as the currency, emphasizing the major factors driving this process and its underlying mechanisms. Likewise, it is proposed that studies aiming at characterizing optimal patterns of plant defense should use chemical defense concentration as the currency, and be accompanied by evaluations of the actual resistance against herbivores of the plant parts analyzed, together with the effect on plant fitness. Received 19 February 1999; accepted 28 April 1999.     

Lignoid chemical defenses in the freshwater macrophyte Saururus cernuus

Summary. Chemical defense against herbivores has rarely been investigated for freshwater plants, possibly due to the common misconception that herbivory on aquatic macrophytes is low and would not select for chemical defenses. In previous work, the freshwater angiosperm Saururus cernuus was shown to be a low preference food for omnivorous crayfish despite its high nutrient value and relatively soft texture. We used feeding by the crayfish Procambarus clarkii to guide fractionation of the deterrent lipid-soluble extract of this plant, leading to the identification of seven deterrent lignoid metabolites, (–)-licarin A, (+)-saucernetin, (–)-dihydroguaiaretic acid, (–)-sauriols A and B, (–)-saucerneol, and (–)-saucerneol methyl ether. Lignans have been implicated in terrestrial plant chemical defenses as insect growth inhibitors, insect toxins, nematocides, antibacterial, and antifungal agents. However, these activities have rarely been demonstrated using ecologically relevant methodologies in terrestrial systems, and never before in freshwater systems. The widespread nature of lignans amongst very distantly related plants, along with their rich diversity of molecular structure, suggests that they could play a large role in mediating plant-herbivore interactions. In addition to the lignoid compounds we identified, there were other compounds present in low concentration or unstable compounds that were deterrent, that did not appear to be lignans, but that we were unable to identify. This plant thus appears to be defended by a complex mixture of natural products. Received 6 June 2000; revised 23 August 2000; accepted 2 September 2000     

Chemical defenses, nutritional quality, and structural components in three sponge species: Ircinia felix, I. campana, and Aplysina fulva

Allocating chemical defenses to regions or tissues most at risk for predatory attack may provide protection while simultaneously minimizing associated metabolic costs. Chemical defense allocation patterns were investigated in the aspiculate sponges Ircinia felix, I. campana, and Aplysina fulva collected between July 2005 and April 2006 from J Reef off the coast of Georgia, U.S.A. It was predicted that chemical defenses would be (1) higher in the outermost 2 mm layer of the sponge; (2) positively correlated with tissue nutritional quality; and (3) correlated with structural components such as spongin fibers. Whereas defensive chemicals were concentrated in the outer 2 mm of A. fulva, the Ircinia species had higher concentrations in deeper tissue layers. Furthermore, no significant positive or negative correlation between chemical defenses and nutritional quality or levels of structural components was observed in these sponges. Overall, these results do not support the prediction that predation pressure by fish and large mobile invertebrates significantly impacts chemical defense allocation in these sponges.     

Allocation of herbivory-induced hydroxamic acids in the wild wheat Triticum uniaristatum

Summary. We characterized the induction of hydroxamic acids (Hx) by aphid infestation in the wild wheat Triticum uniaristatum by addressing the following questions: i) Do different leaves have similar responses to aphid damage?, ii) Is the Hx induction localized or systemic?, iii) How long does the induction last?, and iv) Is the degree of damage related to the magnitude of induced Hx? Based on earlier results on this wheat/aphid system (lack of costs of Hx induction) we expected to find the plant exhibiting cost-saving patterns of response to herbivory. Aphid infestation in the primary leaf led to induced levels of Hx, but no differences in Hx levels were found after infestation of the secondary leaf. Induction of Hx was restricted to the infested leaf (primary leaf). Induced Hx levels exhibited by the primary leaf at the end of aphid infestation were not observed 2 days later. Finally, different aphid densities (between 10 and 40 aphids per leaf) did not produce significant differences in Hx levels in infested primary leaves. Characteristics of Hx induction by aphid infestation in T. uniaristatum partially support the expected cost-saving patterns in the allocation of induced defenses. Received 15 January 1997; accepted 7 July 1997.     

Combining optimal defense theory and the evolutionary dilemma model to refine predictions regarding plant invasion

Optimal defense theory posits that plants with limited resources deploy chemical defenses based on the fitness value of different tissues and their probability of attack. However, what constitutes optimal defense depends on the identity of the herbivores involved in the interaction. Generalists, which are not tightly coevolved with their many host plants, are typically deterred by chemical defenses, while coevolved specialists are often attracted to these same chemicals. This imposes an "evolutionary dilemma" in which generalists and specialists exert opposing selection on plant investment in defense, thereby stabilizing defenses at intermediate levels. We used the natural shift in herbivore community composition that typifies many plant invasions to test a novel, combined prediction of optimal defense theory and the evolutionary dilemma model: that the within-plant distribution of defenses reflects both the value of different tissues (i.e., young vs. old leaves) and the relative importance of specialist and generalist herbivores in the community. Using populations of Verbascum thapsus exposed to ambient herbivory in its native range (where specialist and generalist chewing herbivores are prevalent) and its introduced range (where only generalist chewing herbivores are prevalent), we illustrate significant differences in the way iridoid glycosides are distributed among young and old leaves. Importantly, high-quality young leaves are 6.5x more highly defended than old leaves in the introduced range, but only 2x more highly defended in the native range. Additionally, defense levels are tracked by patterns of chewing damage, with damage restricted mostly to low-quality old leaves in the introduced range, but not the native range. Given that whole-plant investment in defense does not differ between ranges, introduced mullein may achieve increased fitness simply by optimizing its within-plant distribution of defense in the absence of certain specialist herbivores.     

Seaweed sex pheromones and their degradation products frequently suppress amphipod feeding but rarely suppress sea urchin feeding

Summary. A diverse group of brown seaweeds produce bouquets of C₁₁ metabolites, some of which act as pheromones that cue gamete release or attract sperm to eggs following release. We demonstrate that these C₁₁ metabolites and their degradation products also frequently and strongly deter feeding by the herbivorous amphipod Ampithoe longimana, but rarely by the herbivorous sea urchin Arbacia punctulata. Across the range of concentrations tested, seven of twelve C₁₁ metabolites or mixtures that we tested deterred feeding by the amphipod, but only two of eleven deterred the sea urchin. For those compounds where we could rigorously contrast the magnitude of deterrence against the amphipod with the magnitude of deterrence against the urchin, the amphipod was deterred significantly more than the urchin by five of six metabolites. Thus, C₁₁ compounds were more frequently and more strongly deterrent to the amphipod than to the sea urchin. These findings for C₁₁ metabolites conflict with previous investigations, where other classes of seaweed chemical defenses have been shown to deter feeding by large mobile herbivores like urchins and fishes but to be relatively ineffective against mesograzers, especially the species of amphipod that we used here. Our results suggest that C₁₁ metabolites are unusual among the known seaweed chemical defenses in that they are especially effective against mesograzers, which often consume seaweed spores, zygotes, and juveniles. The high concentrations of C₁₁ metabolites in brown algal eggs could allow these defenses to be especially important in defending gametes, zygotes, or young sporelings from herbivorous mesograzers. Received 26 February 1998; accepted 9 April 1998.     

Benefits of trenching behavior in the context of an inducible defense

Summary. Many species of insects sabotage the pressurized defense vessels of their host plants prior to feeding. This behavior, however, does not render leaves indefinitely suitable, as some species employing this behavior eventually abandon uneaten portions of sabotaged leaves. In this study, we examined whether and to what degree wild parsnip, Pastinaca sativa, is capable of restoring its pressurized defenses and whether cabbage loopers, Trichoplusa ni, which normally trench parsnip leaflets, benefit from their trenching behavior. The pressurized oil tubes of parsnip leaves are rich in toxic terpenoids and furanocoumarins. A disruption of the integrity of the tubes (via razor blade nicks) in leaflets revealed that that some of their contents were expelled at the break and that some movement of oil from outside the leaflet (i.e., the midvein) occurred, bolstering furanocoumarin levels in the leaflet within minutes. Pressure and chemical content in a leaflet’s oil tubes were also shown to be restored within 24 hours of depressurization. This recovery ability allowed parsnip leaflets to respond to daily depressurizations by mechanical damage for up to at least 5 assaults, cumulatively causing an approximate ten-fold increase in furanocoumarins. Cabbage loopers fed parsnip leaflets that were artificially trenched accumulated twice as much body mass as larvae fed leaflets augmented with furanocoumarins equivalent to the quantity that would be avoided through trenching, indicating that trenching does benefit the herbivore. Although parsnip recovers from trenching rapidly, it does not do so within the time that cabbage loopers consume trenched leaflets