胡杨雌雄株叶片的比较解剖学研究

应用石蜡切片法对胡杨(Populus euphratica Oliv.)雌、雄株叶片进行比较解剖研究,结果表明:(1)胡杨雌、雄株叶片都由表皮、叶肉与叶脉构成,表皮由2层细胞构成复表皮;上、下表皮内均分化出了2～3层栅栏组织细胞,栅栏组织之间有少量海绵组织细胞;主脉维管束通常1个,由木质部、韧皮部、少量形成层及类似禾本...     

榉树叶片解剖构造和叶肉细胞超微结构的观察

对取自不同季节的榉树(Zelkova schneideriana hand.-Mazz.)叶的解剖构造和叶肉细胞超微结构进行了观察。榉树叶表面被密集锥状单细胞毛,幼叶则还有头状腺毛。气孔多分布在叶下表面,为不规则型。上表皮细胞外壁具有明显的角质层加厚,局部上表皮由2层细胞组成。叶肉细胞分化为栅栏组织和海绵组织;栅栏组织1—2层细胞;海绵组织发达,常与气孔相连。叶肉有含晶细胞存在。叶脉的维管束鞘明显,是厚壁细胞。成熟叶肉细胞细胞器丰富,有8-9个周壁叶绿体,为巨大的长椭圆形,基粒片层清晰,有淀粉粒,少量脂滴。叶片开始转变颜色时,叶绿体由长椭圆形变得趋于圆形;基粒片层逐渐变得杂乱至模糊不清;脂滴数目增多。秋季叶色不同的榉树叶片解剖构造存在一定差异。     

桂花叶解剖结构的研究

通过石蜡切片法制片,光学显微镜观察,研究了桂花叶的形态解剖结构。结果表明,桂花叶为典型的异面叶,表皮由1层排列紧密的形状不规则的表皮细胞构成,细胞外壁角质膜较厚,气孔只分布于下表皮。表皮上被有腺毛,腺毛及其周围排列整齐的表皮细胞形成环式结构。叶肉组织发达,栅栏组织由2～4层排列整齐的圆柱形细胞构成。叶肉中分布有少分枝或不分枝的石细胞,其细胞长轴与表皮垂直。叶脉结构较为复杂,主脉发达。上述特征与复杂多变的环境条件相适应。     

地黄叶和茎的解剖学及组织化学研究

采用解剖学和组织化学方法对地黄叶和茎的显微结构以及梓醇、多糖的分布进行观察研究,以明确梓醇和多糖在地黄叶和茎中的分布特征。结果显示:(1)地黄叶的上、下表皮均分布有腺毛和非腺毛,腺毛都属于头状腺毛,包括长柄和短柄的头状腺毛,两类腺毛的分泌物化学成分主要是黄酮和多糖;叶的上、下表皮上都分布有无规则型气孔,下表皮的气孔密度比上表皮的大,但气孔指数相差不大;栅栏组织由2～3层薄壁细胞构成,排列紧密,海绵组织薄壁细胞形状无规则,细胞间隙大。(2)组织化学研究表明,海绵组织中黄斑样的薄壁细胞是梓醇和多糖的贮存场所,这类薄壁细胞在叶片边缘的齿末端处最为集中,茎的皮层、韧皮部和木质部的薄壁细胞也都是梓醇和多糖的贮存场所。     

阿月浑子与中国黄连木叶形态结构特征研究

采用指甲油印迹法、石蜡切片观察、电镜观察的方法对阿月浑子与中国黄连木叶片解剖结构、气孔分布与形态特征进行观测分析。结果显示,阿月浑子叶上表皮气孔分布密度为132～P168个/cm2,且气孔多出现在主脉附近,下表皮气孔数从基部到叶尖逐渐减少,密度为112～P357个/cm2;中国黄连木上表皮气孔极少,仅出现在主脉处,下表皮气孔分布状况与阿月浑子相近,密度为164～P377个/cm2;两树种均分布有巨型气孔和气孔群、表皮细胞角质层发达、少量非腺毛主要分布在叶缘,其次在主脉处,叶面有蜡质分布,而且阿月浑子的蜡质明显比中国黄连木多;阿月浑子有4～P5层栅栏组织细胞,几乎没有海绵组织,中国黄连木叶肉具1层栅栏组织,多层海绵组织;阿月浑子主叶脉6～P8个维管束环状排列,在韧皮部中有6～P8个内分泌道,中国黄连木主叶脉3个维管束扇形排列,在韧皮部中有1～P3个内分泌道。对2树种叶形态结构特征与抗旱性的关系进行分析结果表明,阿月浑子具有旱生植物叶片结构特点,其水分利用方式属耗水型,对我国北方气候表现出一定的适应性。     

越冬期遮阴条件下3个不同秋眠型紫花苜蓿品种叶片解剖结构与其光生态适应性

在我国南北气候过渡地区, 采用遮阴试验和石蜡切片法, 研究越冬期不同光强对3个不同秋眠型紫花苜蓿(Medicago sativa)品种(‘维多利亚’、 ‘巨人201’和‘游客’)叶片解剖结构的影响。结果表明: 随着光强减弱, 各紫花苜蓿品种表皮结构中上、下表皮角质层厚度, 气孔密度和气孔开度明显下降; 上、下表皮厚度呈上升趋势。随着遮阴强度增加, 叶肉组织中海绵组织细胞宽度显著上升, 栅栏组织厚度、栅栏组织细胞层数、栅栏组织厚度/海绵组织厚度显著下降; 品种间海绵组织厚度和栅栏组织细胞宽度变化趋势不一致。叶片结构整体特征中叶片厚度、叶肉厚度、中脉厚度、组织结构紧密度随光强减弱而显著下降, 组织结构疏松度明显上升, 叶脉突起度变化不明显。品种间各叶片解剖性状变幅及可塑性指数具有明显的差异, 表明其对弱光适应方式不同。Pearson相关分析表明, 各紫花苜蓿品种叶片气孔密度、栅栏组织厚度、叶肉厚度、叶片厚度及栅栏组织厚度/海绵组织厚度与光强呈显著正相关, 可能是紫花苜蓿叶片解剖结构光强敏感特征参数, 其中, ‘维多利亚’叶片敏感特征参数与光强相关程度较低, 与光强相关的性状较少。综合各项分析结果, 初步确定越冬期紫花苜蓿耐阴性与其秋眠性相关, 半秋眠型品种‘维多利亚’ >秋眠型品种‘巨人201’≥非秋眠型品种‘游客’。     

风毛菊属3种植物叶的解剖结构比较

采用石蜡切片法对分布于祁连山海拔5 000 m左右流石滩上菊科风毛菊属水母雪兔子（Saussurea medusa Maxim）、鼠曲雪兔子（Saussurea gnaphalodes (Royle) Sch.）、红叶雪兔子（Saussurea paxiana Diels.）3种植物叶片的解剖结构进行了比较研究,结果表明：叶片表皮细胞均为单层,上下表皮都有气孔分布,气孔不下陷;角质层较厚,叶表面均被单列细胞的表皮毛。3种植物均为异面叶;叶肉栅栏组织较发达,通常由2～3层细胞组成,但栅栏组织细胞排列较疏松;海绵组织存在大量的细胞间隙;叶肉中通气组织发达,且均有不规则裂生性气腔。叶脉维管束中韧皮部都具有异细胞存在。这些共同特征是3种植物对高山地区缺氧、低温、强辐射等自然条件长期适应的结果。但是,3种植物在叶片的外部形态特征、叶肉栅栏组织细胞的特点、维管束发育程度、内分泌结构、不规则裂生气腔等方面又存在明显的不同,表现出3种植物对环境的适应也是存在差异的。     

中国假鹰爪属和皂帽花属植物叶的形态结构及其分类学意义

利用扫描电镜技术、叶片离析法和石蜡切片法研究了假鹰爪属Desmos 4种植物和皂帽花属Dasy-maschalon 3种植物叶片的形态结构。结果表明：假鹰爪属植物叶片近轴面表皮具大型球状含晶簇细胞和不含晶簇的表皮细胞两种类型,远轴面表皮细胞均具一较小的晶簇;叶肉组织明显分化为栅栏组织细胞和海绵组织细胞,油细胞分布于第2层的栅栏组织和海绵组织内,单位毫米叶宽油细胞数为4～6个;主脉维管组织被薄壁细胞分隔成束状。皂帽花属植物叶片近轴面表皮细胞形状相同,均具一晶簇,远轴面表皮细胞的晶簇和近轴面表皮细胞的晶簇相似;靠近上、下表皮的叶肉组织均分化为栅栏组织细胞,在两层栅栏组织细胞之间分化为一至几层海绵组织细胞,油细胞分布于海绵组织内,单位毫米叶宽油细胞数为2～3个;主脉维管组织形成连续的环状。由此可见两属叶的结构具有明显的差异,因而支持假鹰爪属和皂帽花属为两个独立属的观点。     

两种忍冬属植物叶表皮扫描电镜观察及其生态适应性的研究

利用扫描电镜观察2种忍冬属植物的叶表皮形态特征,观察指标包括气孔器、表皮毛、表皮细胞等。扫描电镜观察结果表明:气孔仅分布在下表皮,气孔器散生并为无规则型;下表皮密被表皮毛(包括腺状毛和非腺毛);上表皮细胞的垂周壁呈沟槽状下陷;同时在下表皮还分布有瘤状的草酸钙簇晶。为进一步研究这2种忍冬属植物叶的形态解剖特征,通过光学显微镜观察发现,叶均为异面叶,栅栏组织发达,细胞长柱形;海绵组织细胞类圆形,排列紧密;叶脉机械组织发达。以上这些构造特征,都有利于减少植物体水分的散失,以适应我国西南岩溶区干旱环境特征。通过进一步分析电镜观察结果,发现在忍冬叶的上表皮中脉处有呈线状分布的腺状表皮毛,而华南忍冬叶的上表皮则无表皮毛分布,这一微形态特征可以作为二者种间特异性差别,从而为二者的合理区分提供一定的依据。     

番荔枝科蚁花属和澄广花属叶的比较解剖学研究

利用扫描电镜技术,叶片离析方法和石蜡切片法对蚁花属1种和澄广花属9种植物叶的形态结构进行比较研究。结果表明,两属植物有许多相似之处,但又有以下一些显著不同;蚁花属植物叶表皮细胞均具一晶族,叶肉组织中具1-2层栅栏组织细胞,油细胞均匀分布在栅栏组织和海绵组织中,栅栏组织在主脉处不连续,而澄广花属植物叶的表皮细胞内具一单斜晶,叶肉组织中具1层栅栏组织细胞,油细胞仅分布在海绵组织中,栅栏组织在主脉处连续,结果为蚁花属和澄广花属的分类学处理提供了新证据。     

Leaf anatomy of Desmos and Dasymaschalon (Annonaceae) from China in relation to taxonomic significance

Leaf morphology in four species of Desmos and three species of Dasymaschalon was comparatively studied using scanning electron microscopy (SEM) together with epidermal maceration and paraffin methods. The results showed that there were some remarkable foliar anatomical differences between Desmos and Dasymaschalon. In leaves of Desmos, some of the adaxial epidermal cells were enlarged into globose cells each containing one large cluster crystal, while other epidermal cells were normal without any crystal, and in abaxial epidermis each cell contained one smaller cluster crystal. The leaf structure was typically bifacial, and the mesophyll cells were differentiated into palisade tissue and spongy tissue. Oil cells were distributed in the second layer of palisade and the whole spongy tissue, and the number of oil cells per mm leaf width ranged from 4 to 6. The vascular tissue in the midrib was separated into bundles by parenchyma cells. In leaves of Dasymaschalon, all the adaxial epidermal cells contained one cluster crystal, and the crystal size was similar to that of thecrystals in abaxial epidermal cells. The leaf structure was more or less isobilateral. Oil cells were distributed only in the spongy tissue between the two layers of the palisade, and the number of oil cells per mm leaf width ranged from 2 to 3. The vascular tissue in the midrib formed a continuous circle. It is clear that the anatomical differences between Desmos and Dasymaschalon are remarkable, supporting the treatment of Desmos and Dasymaschalon as two independent genera.     

Patterns of cell elongation in the determination of the final shape in galls of Baccharopelma dracunculifoliae (Psyllidae) on Baccharis dracunculifolia DC (Asteraceae)

Cell redifferentiation, division, and elongation are recurrent processes, which occur during gall development, and are dependent on the cellulose microfibrils reorientation. We hypothesized that changes in the microfibrils orientation from non-galled tissues to galled ones occur and determine the final gall shape. This determination is caused by a new tissue zonation, its hyperplasia, and relative cell hypertrophy. The impact of the insect’s activity on these patterns of cell development was herein tested in Baccharopelma dracunculifoliae—Baccharis dracunculifolia system. In this system, the microfibrils are oriented perpendicularly to the longest cell axis in elongated cells and randomly in isodiametric ones, either in non-galled or in galled tissues. The isodiametric cells of the abaxial epidermis in non-galled tissues divided and elongated periclinally, forming the outer gall epidermis. The anticlinally elongated cells of the abaxial palisade layer and the isodiametric cells of the spongy parenchyma originated the gall outer cortex with hypertrophied and periclinally elongated cells. The anticlinally elongated cells of the adaxial palisade layer originated the inner cortex with hypertrophied and periclinally elongated cells in young and mature galls and isodiametric cells in senescent galls. The isodiametric cells of the adaxial epidermis elongated periclinally in the inner gall epidermis. The current investigation demonstrates the role of cellulose microfibril reorientation for gall development. Once many factors other than this reorientation act on gall development, it should be interesting to check the possible relationship of the new cell elongation patterns with the pectic composition of the cell walls.     

Leaf and green stem anatomy of the drought deciduous Mediterranean shrub Calicotome villosa (Poiret) Link. (Leguminosae)

Light and scanning electron microscopy were used to study leaf and stem fine structure of the drought deciduous green-stemmed Mediterranean shrub Calicotome villosa (Poiret) Link. (Leguminosae). Each leaf consists of three small obovate leaflets with abundant but small (16 μm length) anomocytic stomata on both surfaces. Adaxial surface exhibits more than double stomatal density (440±8 mm⁻²) than the abaxial one (185±4 mm⁻²). T-shaped trichomes (36±3 mm⁻²) are present only on the abaxial leaf surface. Leaves are unifacial, furnished with palisade parenchyma on both sides. The stem is characterized by raised ridges and grooves. Beneath the one-cell-layered epidermis sclerenchyma is found on ridges, whereas stomata and palisade chlorenchyma are found in grooves. Hairs are abundant, especially in grooves. Stem and leaf palisade chlorenchymas are structurally similar. According to these data, photosynthesis could be efficiently supported by the stem.     

泽米科植物羽片脉序和解剖学及其系统学意义

研究了苏铁目泽米科Zamiaceae 2亚科的所有4族(Stevenson系统, 1992)共10种代表植物的羽片脉序及解剖学特征,结果显示泽米科羽片脉序为二歧分叉的平行脉,无中脉。小刺双子铁Dioon spinulosum、大头非洲铁Encephalartos friderici-guilielmii和摩尔大泽米Macrozamia moorei等的平行脉末端以不同的形式互相连接,而鳞木铁Lepidozamia peroffskyana、粗壮角果铁Ceratozamia mexicana var. robusta、竹叶角果铁C. hildae、佛州泽米Zamia　floridana、柔叶泽米Z. debilis、鳞秕泽米Z. furfuracea和短尖泽米Z. muricata等的平行脉末端不连接而直达叶缘,其中鳞木铁、粗壮角果铁和竹叶角果铁的脉达叶缘后逐渐消失。羽片的横切面结构通常由表皮、下皮厚壁细胞和叶肉组成,表皮层包括上、下表皮各一层,叶肉可能同时分化出近上表面的栅栏组织和近下表面的栅栏组织,或仅有近上表面的栅栏组织分化,或无栅栏组织分化而完全为海绵组织。然而,泽米科没有典型的海绵组织和传输组织分化。小刺双子铁、大头非洲铁、鳞叶木铁和摩尔大泽米的羽片具有粘液道而无工字厚壁组织,在小刺双子铁中粘液道与维管束对生,在另3种中则与维管束轮生;但粗壮角果铁、竹叶角果铁、佛州泽米、柔叶泽米、鳞秕泽米和短尖泽米的羽片则具有工字厚壁组织而没有粘液道,其中粗壮角果铁和竹叶角果铁的羽片工字厚壁组织仅与上表皮相连,而佛州泽米、柔叶泽米、鳞秕泽米和短尖泽米的羽片工字厚壁组织与上、下表皮都相连。羽片脉序和解剖学特征支持Stevenson将泽米铁科分为两亚科的观点。     

Ontogenetic differences in mesophyll structure and chlorophyll distribution in Eucalyptus globulus ssp. globulus

Mesophyll structure has been associated with the photosynthetic performance of leaves via the regulation of internal light and CO(2) profiles. Differences in mesophyll structure and chlorophyll distribution within three ontogenetically different leaf types of Eucalyptus globulus ssp. globulus were investigated. Juvenile leaves are blue-grey in color, dorsiventral (adaxial palisade layer only), hypostomatous, and approximately horizontal in orientation. In contrast, adult leaves are dark green in color, isobilateral (adaxial and abaxial palisade), amphistomatous, and nearly vertical in orientation. The transitional leaf type has structural features that appear intermediate between the juvenile and adult leaves. The ratio of mesophyll cell surface area per unit leaf surface area (A(mes)/A) of juvenile leaves was maximum at the base of a single, adaxial palisade layer and declined through the spongy mesophyll. Chlorophyll a + b content showed a coincident pattern, while the chlorophyll a:b ratio declined linearly from the adaxial to abaxial epidermis. In comparison, the mesophyll of adult leaves had a bimodal distribution of A(mes)/A, with maxima occurring beneath both the adaxial and abaxial surfaces within the first layer of multiple palisade layers. The distribution of chlorophyll a + b content had a similar pattern, although the maximum ratio of chlorophyll a:b occurred immediately beneath the adaxial and abaxial epidermis. The matching distributions of A(mes)/A and chlorophyll provide further evidence that mesophyll structure may act to influence photosynthetic performance. These changes in internal leaf structure at different life stages of E. globulus may be an adaptation for increased xeromorphy under increasing light exposure experienced from the seedling to adult tree, similar to the characteristics reported for different species according to sunlight exposure and water availability within their native habitats.     

豆蔻属(姜科)植物叶的比较解剖学研究

采用石蜡切片法对豆蔻属12个种的叶片横切面结构进行比较解剖学研究。结果显示:豆蔻属植物叶片表皮层由1层表皮细胞组成,在无下皮层的种中,近轴面的表皮细胞通常体积较大,而在有下皮层的种中,表皮层通常较薄,细胞壁增厚;下皮层的结构具有多样性;叶肉中栅栏组织通常1~2层,海绵组织2~6层细胞,常含丹宁,所观察的大部分种含有方晶或砂晶;中脉韧皮部极端维管束形态多样,维管束系统II通常缺失,未发现同时具有4个系统的种。此外,对叶缘的形态特点进行了概括性描述,并结合前人的研究资料讨论了叶解剖结构的系统学意义。     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

Anatomy and uses of the mature leaves of three species of Sabal (Arecaceae) of the Yucatan, México

This paper describes the leaf anatomy of Sabal mauritiiformis (Karst.) Griseb. & H. Wendl., Sabal mexicana Mart. and Sabal yapa Wright ex Becc., three of the four most representative species of the Yucatán Península, in Mexico. These species are locally used: in the roofing of traditional homes, as food (fruits and apical buds), and in the production of hats, brooms and handicrafts. Leaf samples were collected in secondary growth of lower montane rainforest in the state of Quintana Roo and in two home gardens in the state of Yucatán. Herbarium samples were obtained, and samples of blade and petiole were fixed in formaline-acetic acid-alcohol. Cross incisions were made on the blade and petiole, and were dyed with safranin and toluidine blue O. The results show that S. mauritiiformis and S. yapa are morphologically alike: both are tall, slim palm trees; the leaf in S. mauritiiformis is a shorter palm-like structure compared with the other two species. The shape of the main nerve, as seen in cross section, is rectangular in the three species. The hastula in the three species is acuminate and adaxial. The foliar anatomic structure is similar in the three species, although there are some differences. The adaxial an abaxial epidermis of the blade consist of one layer and, superficially, the anticlinal walls are straight; the stomata are intercostal, of the tetracytic type, present on both surfaces in S. mexicana and S. yapa and only on the abaxial surface on S. mauritiiformis. The hypodermis is one layer thick in S. yapa and in S. mexicana and two layers thick in S. mauritiiformis. In the three species the palisade parenchyma consists of several undefined strata as the cells are similar-in shape and size--to the cells in the spongy parenchyma, so there is no marked difference between these strata and the spongy parenchyma seems almost continuous. Both fibrous and vascular bundles are distributed between the hypodermis and the palisade parenchyma; the fiber bundles can be found towards the abaxial surface while the vascular and fiber bundles are located towards the adaxial surface. The fibers, in the three species, are elongated, with the pointed tips, undivided and unseptated. One to three wide vessels of metaxilem can be seen in the vascular bundles, those in S. yapa being the widest in diameter. The vascular bundles are surrounded by thick fiber sheaths which come in pairs. The anatomic structure of the petiole is similar to that of the blade, and is characterized by the many vascular and fiber bundles dispersed in the parenchymatous tissue, and which are very resistant. The histological structure of the blade and petiole reflects strength and flexibility, qualities which make these plants adequate in the construction of roofs for rural housing and other buildings.     

Micromorpho-Anatomical Examination of 2,4-D Phytotoxicity in Grapevine (Vitis vinifera L.) Leaves

The anatomical and micro-morphological alterations as induced by the auxinic herbicide, 2,4-D (2,4-dichlorophenoxy acetic acid) have not yet been elucidated for a commercially important fruit crop such as grapevine despite its super sensitivity to 2,4-D. Light and scanning electron microscopy techniques were employed to examine 2,4-D induced internal and external structural abnormalities in Merlot grapevines (Vitis vinifera L.). Healthy leaves were dorsiventrally flattened with well developed patterns of cellular structure and composition involving adaxial palisade parenchyma and abaxial spongy mesophyll. Dorsiventral variations in epidermal features involved large epidermal cells on the adaxial surface, and trichomes and stomata with turgid elliptical guard cells on the abaxial surface. The 2,4-D injured leaves were small and enated; the veins were fasciated with rugose bands of lamina existing between fasciated veins. The epidermal cells aggregated instead of being positioned coplanar to the epidermal plane. The adaxial elongated palisade parenchyma cells were transformed into an ovoid shape with intercellular spaces. An extensive development of replacement tissues took place on the abaxial surface wherein the stomata became roundish and were either raised or sunken with collapsed and cracked guard cells that developed abnormal outer stomatal ledges. These abnormalities are expected to severely perturb the vital functions of photosynthesis and transpiration ultimately leading to vine death attributable, at least in part, to the injured leaves.